Signalling plays a key role regulating the intensity of animal contests. In this study, we evaluate the role of two (potential) chromatic signals in determining the outcome of male-male combats in Podarcis muralis lizards from a population showing polymorphic ventral coloration, with discrete white, yellow and orange morphs. As in other lacertids, males of this species display a row of ventrolateral ultraviolet (UV)- blue in their outer ventral scales (OVS) with a mean peak reflectance of 372.35nm. Previous studies have suggested that some chromatic variables of these patches may signal individual fighting ability and/or physical condition. In order to test this possibility, we staged combats between 60 adult lizards (20 individuals/morph), where each lizard confronted rivals from each morph in a tournament with a balanced design. Intruder/resident condition was controlled in order to avoid its strong effect in determining contest outcome. A general dominance/fighting ability ranking was calculated using the Bradley-Terry model, and used to explore the general relationship between morphs, chromatic measurements of the UV patches, and fighting ability. Unexpectedly, we did not find an effect of the UVblue patches on contest outcome, but we found a morph-specific bias in fighting ability: orange males lose more combats with other morphs than yellow and white males.

The role of male coloration in the outcome of staged contests in the European common wall lizard (Podarcis muralis). - Behaviour, 153 (5): 607-631.

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Colour signals play a key role in regulating the intensity and outcome of animal contests. Males of the common wall lizard (Podarcis muralis) show conspicuous ventrolateral ultraviolet (UV)-blue and black patches. In addition, some populations express a striking ventral colour polymorphism (i.e., discrete orange, white and yellow morphs). In this study, we set out to evaluate the potential signalling function of these colour patches by staging pairwise combats between 60 size-matched adult lizards (20 per morph). Combats were held in a neutral arena, with each lizard facing rivals from the three morphs in a tournament with a balanced design. We then calculated a fighting ability ranking using the Bradley–Terry model, and used it to explore whether ventral colour morph, the size of UV-blue and black patches or the spectral characteristics of UV-blue patches (i.e., brightness, hue, chroma) are good predictors of fighting ability. We did not find an effect of the UV-blue patches on contest outcome, but the size of black patches emerged as a good predictor of fighting ability. We also found that winners were more aggressive when facing rivals with black patches of similar size, suggesting that black patches play a role in rival assessment and fighting rules. Finally, we found that orange males lost fights against heteromorphic males more often than yellow or white males. In light of these results, we discuss the potential signalling function of ventrolateral and ventral colour patches in mediating agonistic encounters in this species.

The Italian lizard (Podarcis sicula) is the most diffused reptile in Italy, but it is also present in other European countries. This lizard belongs to the Lacertidae family, lives near walls, slants and along the borders of the paths; its diet includes bugs and aracnids. No data are so far available in literature about the three-dimensional morphology of the tongue of Podarcis sicula, therefore the aim of the present paper was to study by scanning electron and light microscopy the three-dimensional characteristics of the dorsal lingual surface and moreover the presence of chemosensory receptors like the taste buds in the oral cavity. Our results demonstrate that the Podarcis sicula tongue is a triangular muscular membranous organ, dorsoventrally flattened and that three different areas can be observed: a bifid apex, a body and a root. No papillae were observed in the apex, characterized by a flattened mucosa and by two deep median pouches. In the body cylindrical papillae with a flat surface are present, aborally gradually substituted by imbricated papillae. Foliate-like papillae were observed in the lateral parts of the tongue body. No sensory structures were showed on the lingual dorsal surface, while they were numerous in the oral cavity, particularly on the gingival epithelium. The light microscopy shows, on the dorsal surface, a stratified pavimentous not keratinized epithelium, conversely keratinized along the ventral surface. Many caliciform cells on the lateral parts of the papillae, deputed to the secretion of mucus, were also observed. Therefore, the results obtained in this paper could give a contribution to the knowledge of the tongue anatomy in a species widely diffused in different European countries and could be of help for clinical purposes in reptiles.

The Herpetofauna of the Gaza Strip with Particular Emphasis on the Vicinity of Wadi Gaza. - The Islamic University Journal (Series of Natural Studies and Engineering), 15 (1): 111-135.

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The reptiles and amphibians of the Gaza Strip and Wadi Gaza were
surveyed during a period of two years (2002 – 2004). A total number of 21
herpetofaunistic species (2 turtles, 8 lizards, 8 snakes and 3 anurans)
belonging to three orders and 15 families was encountered. The species
described were all resident and were mostly found throughout the year. The
diversity of terrestrial and aquatic ecosystems in the study area encouraged the
occurrence of the species. However, the ever-increasing human impact on the
existing natural resources in the Gaza Strip has threatened the ecology of
wildlife, where the populations of frogs and many reptilian species are
declining in an alarming fashion. The results reinforce the necessity of longterm
inventories in order to understand the ecology and the dynamics of
herpetofaunistic and other wildlife communities in the study area. Finally, the
authors recommend improving cooperation of different parties to enhance the
public awareness among the Palestinians and to implement environmental
laws and legislation to conserve the sensitive and rare species of herpetofauna.

The cranial musculature of species of ten families of lizards (Gekkonidae, Teiidae, Gymnophthalmidae, Anguidae, Scincidae, Lacertidae, Tropiduridae, Liolaemidae, Leiosauridae, and Polychrotidae) was analyzed. Using 93 myological cranial characters, a cladistic analysis was performed. To root the trees, data of Sphenodon were added to the matrix. The cladistic analysis yielded 129 equally parsimonious trees with a fit of 426.0 (39%) and 467 steps. In the consensus tree we observed only five nodes congruent with traditional phylogenetic hypothesis of the groups analyzed: teiids + gymnophthalmids (node 128); geckos (node 136); liolaemids (node 116); genus Liolaemus (node 115); and genus Phymaturus (node 92). Thus in these groups, phylogeny seems to be better predictor of muscle morphology than ecology or diet. They are many problematic groups whose monophyly requires further analysis (tropidurids, polychrotids, and scincids). In no case we were able to recognize differences that could be attributed to functional features.

This paper describes the morphological characteristics of a new species of Haemogregarina Danilewsky, 1885, from naturally infected Acanthodactylus boskianus (Daudin) in Egypt, based on both blood and tissue stages. The prevalence of infection was found to be 66% (33/50). The blood stages were found to be confined to the erythrocytes and could be differentiated into small and large forms. The small form (trophozoites) measures 8 × 3.5 μm, whereas the large form (mature gamonts) measures 12 × 5 μm. The tissue stages were observed only in the liver. Meronts occurred within parasitophorous vacuoles in hepatocytes and ranged from mononucleate forms to mature forms that contained c.10–15 merozoites.

To date, three species of Isospora Schneider, 1881 have been described from lizards of the genus Acanthodactylus Wiegmann. Two of these, although representing separate species parasitizing two different hosts, Acanthodactylus boskianus Daudin in Egypt and A. schmidti Haas in Saudi Arabia, were described under the name Isospora acanthodactyli. The third species is Isospora abdallahi Modrý, Koudela et Slapeta, 1998 from A. boskianus in Egypt. In the present study, Isospora alyousifi nom. n. is proposed to accommodate Isospora acanthodactyli Alyousif et Al-Shawa, 1997 (homonym of I. acanthodactyli Sakran, Fayed, El-Toukhy et Abdel-Gawad, 1994) and its redescription based on newly collected material is provided.

Haemogregarina damiettae was previously described form Acanthodactylus boskianus in Egypt, and the description was base only on the blood stages with no data on the tissue stages. In the present study, both blood and tissue stages were encountered. The prevalence of infection was found to be 60 % (30/50). The blood stages were found to be confined to the erythrocytes and none in the leucocytes. The erythrocytic stages could be differentiated into small and large forms. The small form, trophozoite, measured 10 × 3 μm, while the large form, mature gamontocyte, measured 18 × 5 μm. The cytoplasm of the mature gamontocyte was finely granular and faintly stained with Giemsa, while the nucleus was relatively large and occupied about one third of the body length and was formed of a network of chromatin filaments. Infected erythrocytes were hypertrophied and their nuclei either longitudinally stretched or split into two fragments. The tissue stages were observed in the lungs and hearts of the infected lizards. The merogonic stages were evident as multinucleated ovoid to subspherical bodies enclosed by parasitophorous vacuoles. Mature meronts measured 16 × 12 μm and estimated to produce 15–25 merozoites.

Chemical treatments still constitute the main method used to control desert locust during the invasion phases. In addition to their relatively high costs, these pesticides are not target specific in the majority of the cases; they can have negative effects on the biotic and abiotic components of the ecosystem. The goal of our study, conducted under natural conditions, was the assessment of the ecological effect of ethyl chlorpyrifos and fenitrothion, the most widely used organophosphate pesticides in desert locust control in Niger. To assess the impact of pesticides on the relative abundance of lacertid lizard Acanthodactylus boskianus and Acanthodactylus sp before and after treatments, we used the method of pedestrian transect. The analysis of the variance was used to carry out a statistical comparison based on BACI (Before-After-Control-Impact) method. The experiments were carried out in a randomized complete block design with three replications. The experimental plots had an area of 16 ha each, the treatments consisted of ethyl chlorpyrifos at the rate of 225 ga.i./ha; fenitrothion at the rate of 450 ga.i./ha. Unsprayed plots were used as controls. These rates are those recommended by the Food and Agriculture Organisation of the United Nations (FAO) to control desert locust. Chemical treatments were carried out in total cover using a battery driven hand-held spinning disk sprayer (Micro-Ulva®), in accordance with the technical directives of the FAO for ultra-low volume applications. The field trial was carried out in the valley of Tafidet located in the eastern side of Aïr Mountains in Niger, from June to December of the years 2004 to 2006. Ineach experimental plot, two lines of 300 m length and 10 m width were used as area of counting. These lines of counting skirted one of the diagonals of each plot. The results demonstrate the noxious effect of ethyl chlorpyrifos and fenitrothion (organophosphate pesticides) on lizard. We recorded the first decrease of population at 9 days after treatment. The first decrease of the lizard populations was noted in the phase I (1-12 DAT). We also recorded the first dead or moribund lizards between 9 and 21 days after chemical treatment. The number of death or moribund lizards ranged from 0, 33±0.57 to 4±1 in the plots treated with chlorpyrifos and fenitrothion respectively. The optimum of the efficacy of the pesticides was noted in phase II (15-30 DAT) it was decreased from 76% to 98% with chlorpyrifos and fenitrothion. It was also observed; the chemicals had a delayed effect on the lizard. In the last phase (45-60 days after the treatment) no death or moribund lizards is noted in all plots (treated and untreated).

The patterns of seasonal and daily activities of two parthenogenetic species of lizards (Darevskia armeniaca, D. unisexualis) sharing the same territories in the zone of hybridization in central region of Armenia demonstrate ecologically similar potential niches.

Parthenogenetic lizards of the genus Darevskia as an evolutionary model.
Several parthenogenetic lineages occur within the lizards of the genus Darevskia (Sauria: Lacertidae) which are endemic to southern Transcaucasus. High level of heterozygosity, cause by thein hybrid origin, is one of the crucial aspects of thein evolutionary potential, as well as the asexual reproduction. Heterosis on one side is in the opposition to the outbreeding depression and genetic uniformity of the clones on the other side. Aim of this work is to evaluace if these aspects influence viability of parthenogenetic species and differ them from the sexual ones. We chose the amount of asymmetries as a measure of developmental instability, which we studied on three meristic characters. We also evaluated the pattern of asymmetries in lateral blue spots, which are of signaling importace in lacertid lizards. Our results suggest that there isn`t significant difference between parthenogenetic and sexual species in developmental stability, but the sexual ones are more sensitive to population changes. Absence of males may have perhaps the greatest influence on coloration, resulting in loss of symmetry in the blue spots.

Raillietiella morenoi sp. n., a new cephalobae- nid pentastomid found in the lungs of Gallotia atlantica collected in the Canary Islands, is described. The new species belongs to the sharp-tipped posterior-hook type. The annulus number, morphology, and dimensions of copulatory spicules and the dimensions of anterior and posterior hooks separate Raillietiella sp. n. from the other raillietiedid sharp-tipped posterior-hook species of small lizard parasites in Africa. The host character of endemic protected species of G. atlantica in Alegranza Island posed great difficulty in obtaining more parasite materials. However, more studies are required to state the variability of this species and its possible distribution in other species in the Canary Islands, as well as in other Gallotia spp.

In the course of the faunistic inventory in Wadi Ramm Protected Area, a total of 34 species of reptiles representing
nine families (Gekkonidae, Chamaeleonidae, Agamidae, Lacertidae, Scincidae, Varanidae, Leptotyphlopidae,
Colubridae, and Viperidae) were recorded from different habitats in Wadi Ramm and its closest vicinity.
Three species (Lacerta cf. kulzeri, Chamaeleo chamaeleon, and Ablepharus rueppellii) are considered to represent
relicts from earlier, more humid periods. Habitat preferences for collected species are included.

This study reports on the seasonal, sex, and age differences in the diet composition of a population of the lacertid lizard Podarcis milensis inhabiting an arid sand dune on Milos Island (Aegean Archipelago, Greece). Stomach contents of 191 animals were analyzed and compared with prey availability data. The most important prey types were Hemiptera, Coleoptera (other than Tenebrionidae), spiders, and ants. Other prey items included plant material and insect larvae, both constantly consumed throughout the year. Differences in the taxonomic composition of the diet were found between males and females and between juveniles and all other classes. The greatest similarity of diet composition between the sexes and ages examined was during summer. The foraging patterns observed are discussed within the context of the specific study system: an insular, low productivity, sand dune ecosystem.

Effects of body size, throat color and residency on the outcome of intrasexual encounters in the males of the lacertid Podarcis milensis: ecological and evolutionary implications. - 6th International Congress of Systematic and Evolutionary Biology, Patras, Greece, 9-16/9/2002. Book of Abstracts: 267.

The reproductive strategy of Podarcis milensis exhibits some peculiarities when
compared with other congeners. Males and females attain sexual maturity at a min-
imum body size of 47 and 42 mm SVL, respectively, both at an age of about one
year. Podarcis milensis has a ver y small clutch size, with a mean of 1.73 and a range
of 1–3 eggs, and produces multiple clutches annually. Both sexes exhibit a pro-
longed reproductive period extending from Januar y to August.

Field body temperatures (Tbs), activity cycles, and preferred body temperatures maintained in a laboratory thermogradient (Tsel) were studied for Podarcis milensis, a small, endemic, lacertid lizard occurring in Milos Archipelago, Greece. P. milensis is active all year round; overall maximum activity level is recorded in spring, and minimum activity in winter. Daily activity patterns range from unimodal (winter) to strongly bimodal (summer). Body temperature of adults of the examined population (n = 188) averages 31.3 °C (range 21.5-38.4 °C, SD 3.27 °C); mean monthly Tbs are grouped together into `seasons`. The species actively thermoregulates, and effectiveness of thermoregulation for the month of August is high, 0.95. The thermoregulatory behavior, microhabitat utilization, and activity cycle of this population are all discussed in the specific context of our study system: the harsh thermal environment of an insular sand dune.

On the distribution and ecology of amphibians and reptiles in the south of the Russian Far East. - Russian Journal of Herpetology, Moscow, 13 13 (2): 101-116.

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The paper reports on the distribution and ecology of amphibians and reptiles in the south of the Russian Far East,
including previously unstudied areas within Khabarovsk Kray, Primorskiy Kray, and Yevreyskaya Autonomous
Oblast’ (209 localities in total) studied in 1998 – 2004.

Adnagulov, E.V. (2017) -

Annonated list of Amphibian and Reptile species of the Far East Russia. - Current Studies in Herpetology, Saratov, Russia, 17 (3-4): 95-123.

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An annotated list of amphibian and reptile species of the Russian Far East is presented with due account
of modern data on the taxa systematics. 13 Amphibian species and 27 reptile ones have been reliably registered
by now, including several random sea species.

O montado, sistema agro-silvo-pastoril de extrema importância para várias espécies de répteis, foi moldado pela ação humana desde tempos ancestrais e a manutenção das suas características seminaturais depende diretamente da continuação da gestão tradicional. A presente investigação teve como principal objetivo a determinação do impacto da gestão do montado, principalmente do pastoreio, no uso do habitat pelas lagartixas Psammodromus algirus, Psammodromus occidentalis e Podarcis virescens. Os resultados atingidos, por meio da realização de testes de hipóteses, apontam para a ausência de impacto dos vários níveis de intensidade de pastoreio no número de indivíduos encontrado em cada cerca, embora, aparentemente, haja uma alteração do comportamento e uso do micro-habitat pelas lagartixas quando expostas às diferentes intensidades. Estas lagartixas mostraram selecionar micro-habitats com uma estrutura simples, sendo que as diferenças entre as espécies se encontram no uso que estas fazem dos elementos que o constituem; Abstract: Habitat Use by Lizards in Agrosilvopastoral Systems of Montado The montado is an extremely important agrosilvopastoral system to several species of reptiles. It has been moulded by the human activity since the primordial times and its characteristics rely directly on the continuity of the traditional ways of management. The present research`s objective has been the measure of the impact inflicted by the management applied on the montado upon the use of the habitat by the species of lizards Psammodromus algirus, Psammodromus occidentalis and Podarcis virescens. The results suggests that there is no impact on the population of the species studied due to the intensity of the grazing, although there has been noticed a change in the behaviour and usage of the micro-habitat by them, when exposed to the different intensity levels. The lizards seem to prefer simple structured micro-habitats being the main difference between the species in the usage that they give to its physical elements.

Afrasiab, S.R. & Mohamad, S.I. & Hossain, R.H. (2013) -

A review of the Lacertini of Iraq in Iraqi collections. - Herpetozoa, Wien, 23 (3/4): 93-100.

In this study, 29 reptile and amphibian species were recorded in 25 diff erent localities in the Sultan Mountains.
Of these, 5 were anurans, 1 was a tortoise, 1 was a turtle, 11 were lizards, and 11 were snakes. A chorotype classifi cation
of the species recorded in the Sultan Mountains is also given.

Two new species of the Ophisops microlepis (Squamata: Lacertidae) complex from northwestern India with a key to Indian Ophisops. - Journal of Natural History, London. Doi: 10.1080/00222933.2018.1436203

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We describe two new species of the lacertid genus Ophisops based on a series of 19 specimens from semi-arid habitats in the states of Gujarat and Rajasthan in northwestern India, provide a description of Ophisops microlepis sensu stricto, and a key to Indian Ophisops. Ophisops pushkarensis sp. nov. and Ophisops kutchensis sp. nov. are allied to Ophisops microlepis and can be diagnosed from all other Indian Ophisops by the fusion of the lower and upper eyelids, their large body size (snout to vent length > 50 mm), and ≥ 50 scales around midbody. They differ from O. microlepis and each other in the number of scales around midbody, the number of dorsal scales, subtle colour pattern differences, as well as uncorrected mitochondrial sequence divergence (6–9%). These are some of the only known endemic reptiles in these semi-arid landscapes and indicate that many other such habitats may harbour endemic biodiversity.

Aim India is dominated by tropical grassy biomes (TGBs), traditionally considered seres or degraded forest, with low diversity relative to the restricted, ancestral wet zone. It is unclear if Indian grasslands and other open habitats are anthropogenically derived or native, old-growth habitats; without a clear timescale of grassland evolution. One way to understand grassland evolution is to study the diversification in taxa restricted to open habitats. We use a dated phylogeny of Ophisops to address questions related to the origin, diversification and inter-relationships of Indian and Saharo-Arabian Ophisops, and ultimately the origin of Indian grasslands and open habitats. Location The Indian subcontinent; the Saharo-Arabian Realm. Methods We generated up to 2736 base pairs of aligned sequence data (one mitochondrial, two nuclear genes) for Indian lacertids and reconstructed phylogenetic relationships using maximum likelihood and Bayesian inference. We use a fossil-calibrated timetree, diversification analyses and ancestral area reconstructions to test the hypotheses of origin and relationships with Saharo-Arabian Ophisops. Results Ophisops is strongly supported as monophyletic, with a basal split into a large-bodied (LBC) and small-bodied clade (SBC). The Saharo-Arabian species are nested within the Indian species in the LBC. Species diversity in Indian Ophisops is grossly underestimated, with 26–47 candidate species. Ophisops began diversifying in the late Oligocene with significant rate shifts in the late Miocene-Pliocene and Pleistocene within the SBC. Main conclusions Our results are consistent with an ancient origin of grassland taxa and TGBs in India. Ophisops is a dramatic example of overlooked cryptic diversity outside forests, with ~30 species where five were known. Ophisops dispersed into India from the Saharo-Arabian Realm in the Oligocene with a back dispersal in the Middle Miocene, a novel biogeographical pattern. Diversification in the SBC of Ophisops increased 8-fold during the global C4 grassland expansion. Indian TGBs are old-growth ecosystems that need urgent conservation attention.

The VIP/VPACR system in the reproductive cycle of male lizard Podarcis sicula. - General and Compoarative Endocrinology, 205: 94-101.

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Starting from the knowledge that in the reproductive period the Vasoactive Intestinal Peptide (VIP) is widely distributed in Podarcis sicula testis, we studied VIP expression and the localization of the neuropeptide and its receptors in the testis of the Italian wall lizard P. sicula in the other phases of its reproductive cycle (summer stasis, autumnal resumption, winter stasis, spring resumption). By Real Time-PCR, we demonstrated that testicular VIP mRNA levels change during the reproductive cycle, showing a cyclic trend with two peaks, one in mid-autumnal resumption and the other in the reproductive period. By in situ hybridization and immunohistochemistry, we demonstrated that both VIP mRNA and protein were widely distributed in the testis in almost all the phases of the cycle, except in the early autumnal resumption. As regards the receptors, the VPAC1R was localized mainly in Leydig cells, while the VPAC2R showed the same distribution of VIP. Our results demonstrate that, differently from mammals, where VIP is present only in nerve fibres innerving the testis, an endotesticular synthesis takes place in the lizard and the VIP synthesis changes throughout the reproductive cycle. Moreover, the VIP/VPAC receptor system distribution observed in germ and somatic cells in various phases of the cycle, and particularly in the autumnal resumption and the reproductive period, strongly suggests its involvement in both spermatogenesis and steroidogenesis. Finally, the wider distribution of VIP in lizards with respect to mammals leads us to hypothesize that during the evolution the synthesis sites have been transferred from the testis to other districts, such as the brain.

Molecular cloning of VIP and distribution of VIP/VPACR system in the testis of Podarcis sicula. - Journal of Experimental Zoology, Part A: Ecological Genetics and Physiology, 321 (6): 334-347.

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Using molecular, biochemical, and cytological tools, we studied the nucleotide and the deduced amino acid sequence of PHI/VIP and the distribution of VIP/VPAC receptor system in the testis of the Italian wall lizard Podarcis sicula to evaluate the involvement of such a neuropeptide in the spermatogenesis control. We demonstrated that (1) Podarcis sicula VIP had a high identity with other vertebrate VIP sequences, (2) differently from mammals, VIP was synthesized directly in the testis, and (3) VIP and its receptor VPAC2 were widely distributed in germ and somatic cells, while the VPAC1R had a distribution limited to Leydig cells. Our results demonstrated that in Podarcis sicula the VIP sequence is highly preserved and that this neuropeptide is involved in lizard spermatogenesis and steroidogenesis.

Aguado de la Paz, S. (2016) -

Thermoregulatory optimization in cold environments: mechanisms in several species of Sauropoda. - Tesis, Repositorio de la Universidad de Oviedo. 191 pp.

Thermal constraints may limit the physiology and behaviour of ectotherms because of the high thermal dependence of metabolic functions. The adaptive mechanisms of thermoregulation and the cost of confronting thermal constraints were studied in the Cyren’s Rock Lizard (Iberolacerta cyreni (Muller and Hellmich, 1937)), a lacertid lizard endemic to mountain areas of central Spain. Semicontinuous monitoring of body temperature (Tb) in the laboratory indicated that the preferred temperature range for this population (Tpref) was lower than those found for most lacertid lizards, and field body temperatures of active animals in summer were even lower than Tpref. Overall these results, together with distribution of field operative temperatures (Te), indicate that I. cyreni is an active and relatively accurate thermoregulator, although limited by thermal constraints in their habitat. Laboratory experiments in contrasting thermal environments showed that even under thermally restricted conditions, lizards achieved their Tpref by modifying their thermoregulatory behaviour, principally through changes in space use, basking time, and body posture. However, these behavioural adjustments to reach the Tpref have associated costs, and lizards spent 80% of their time in thermoregulation when tested under low radiation conditions, which in the wild would limit the scope for other activities and eventually increase predation risk. Our results suggest that thermoregulatory behaviour may play an important role in coping with global climate change, hence predictions of the effects of climate warming on lizards inhabiting cold habitats should take into account the buffering role of behavioural thermoregulation.

The Madeira wall lizard, Teira dugesii selvagensis, is a partially herbivorous species that also predates Cory shearwater chicks, having an impact of 5% on the reproductive success of this seabird at the island of Selvagem Grande. This study aims to characterize and compare the population structure of Madeira wall lizards, as well as its role in the trophic web of various areas of Selvagem Grande. These areas were designated according to the type of relief, size of seabird colonies and vegetation cover, in two different Cory shearwater nesting seasons (egg incubation and offspring feeding period). The abundance of lizards in different areas and in both seasons was estimated using capture-mark-recapture. Lizard droppings were also collected, to identify contents. Samples of species belonging to the Selvagem Grande trophic chain were collected for isotopic analysis. Isotopic results were analyzed using SIAR and SIBER packages. Both the highest abundance of lizards and the highest vegetation cover were found in areas with small seabirds colonies. In spite of that, the proportion of lizard immatures was higher on the South/Southwest facing cliffs, where the largest seabird colonies are located. Herbivory increases with lizard size, and there was a high percentage of vegetable contents (about 65%) in the droppings. The isotopic basis of the trophic chain varied significantly among areas and is probably influenced by island topography and seabird inputs. Comparing signatures between geckos and lizards of the island, allowed the characterization of niches occupied by each species, being the differences justified by their diets. In areas with larger seabird colonies, seabirds or nutrient inputs brought by them were extremely important for the diet of T.d. selvagensis. The lizards with smaller differences in their isotopic niche between seasons were found in the areas with most seabirds, while the highest isotopic niche differences occured where resources were fewer. Isotopic results validate the dropping analysis, indicating consumption of seabirds during the bird offspring feeding period. Results obtained in this study suggest that the pressure of lizards on seabirds colonies is variable along the island. In contrast to what was expected, in areas with more vegetation there was a reduced impact on seabirds, despite higher abundances of lizards. Lizard monitorization should be continued to assess population trends as vegetation recovers after the eradication of rabbit and domestic mouse.

A Contribution on Lacerta media Populations in Agricultural Environments in North West of Iran. - In: Pafilis, P., Kotsakiozi, P. & E.D. Valakos (eds.): 6th International Symposium on the Lacertids of the Mediterranean Basin, 23-27 June 2008, Mythimna, Lesvos, Greece: 11.

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Two species of Lacerta genus are found in Northwest of Iran:
Lacerta media and Lacerta strigata. Lacerta media has wide
distribution range and is an important lizard fauna element in agro-
ecosystems and river closed humid areas. The aim of this study was
to provide reasons for high population size of this species in agro
ecosystems. So, this study was performed in the farming areas in
Meshkinshar, Ardabil province of Iran from 2003 to 2006. Based
on extensive field research, it has been mainly found that Lacerta
media is the most abundant reptile species in mentioned habitats
and it just inhabits areas that are much closed to water. Our results
also showed, animal is hidden by dense grass vegetation cover
especially male that has green color. They use old hallows of mice
and other small vertebrates for hiding and egg lying. In spring they
are seen on cultivated filed boundaries on stones on sunny days. As
specific result, they are found in apple orchards more abundant than
other cultivated fields. It seems that permanent food accessibility
(insects) and fewer predators are most reasons. Finally, Lacerta
media is interesting species to study from a conservational point of
view because of its unusual frequency of occurrence as parallel with
human activities.

A molecular phylogeny of the eastern group of ocellated lizard genus Timon (Sauria: Lacertidae) based on mitochondrial and nuclear DNA sequences - Amphibia-Reptilia 33 (2012): 1-10

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Timon, a small genus of lacertid lizards, includes four species distributed in two separate ranges in the western
and eastern part of the Mediterranean Basin. Phylogenetic relationships between the two groups have not been resolved, and the taxonomic situation of the two subspecies of the eastern representative of the genus, Timon princeps, is not clear. To address these questions, partial DNA sequences of two nuclear (β-fibrinogen intron 7 and C-mos) and three mitochondrial (cytochrome b, 12S rRNA and 16S rRNA) genes were analyzed. Based on the high genetic distance between the two subspecies of T. princeps we promote their taxonomic status to full species, Timon princeps and Timon kurdistanicus.
Divergence time estimates based on other lacertid species suggest that the separation of the green (Lacerta) and ocellated (Timon) lizards took place around 12 My ago, and that the Eastern group underwent speciation around 4-5 my ago, perhaps associated with the uplifting of the Zagros mountains. As expected given this ancient divergence and complex paleogeography, considerable levels of genetic diversity are recovered within both taxa, with geographically close individuals showing very divergent haplotypes.

A variety of different methods are available for investigating the effects of past climate fluctuation on biota. Among them, molecular phylogeography and species distribution modeling approaches have been shown to be especially useful tools to trace past climate induced modifications of species’ geographic distributions. The results of both analytical approaches are here combined to better understand the influences of past climate changes on the fragmented distribution pattern of the lizard genus Iranolacerta that currently occurs in the northwest and central Zagros Mountains of Iran. While Iranolacerta zagrosica is restricted to few localities and cannot be modeled, the potential distribution of Iranolacerta brandtii suggested a large region of unfavorable habitat between northwest and the central Zagros Mountains populations. From the phylogenetic perspective, both species display deep genetic separation likely predating the Pleistocene. However, limited genetic diversity across this divided range of I. brandtii implies that the current separation is recent. Deeper genetic divergences were, nevertheless, uncovered within the northwest population. Since no recent geological events explain these results, the most likely explanation for such a distribution pattern is due to past climate fluctuations. Both sources of evidence suggest that during the early Holocene the northwest and Zagros populations of I. brandtii were connected, and the current pattern was shaped with their disconnection in the early Holocene. Further studies may identify potential glacial refugia for other species in this biodiverse region.

Separate histories in both sides of the Mediterranean: phylogeny and niche evolution of ocellated lizards. - Journal of Biogeography, 43: 1242-1253.

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Aim The ocellated lizards of the genus Timon, comprising six species that are
distributed across continental Asia, Europe and Africa, offer an interesting
model to study the evolution of ecological niches through comparative phyloclimatic
analysis. Our study provides insights into the evolutionary history of
the ocellated lizards and helps to understand the role of climatic niche evolution
during the speciation process.
Location Eastern and western margins of the Mediterranean basin.
Methods A dated molecular phylogeny was estimated based on three mitochondrial
and two nuclear genes. Using multivariate statistics, species distribution
models were developed to characterize the Grinnellian niches of all
species. Subsequently, ancestral environmental niche occupancy of each taxon
was reconstructed using niche occupancy profiles. Niche divergence among
species was quantified by computing multivariate niche overlaps via twodimensional
and n-dimensional approaches.
Results Phylogenetic analysis supports that the ancestor of Timon diverged
into the eastern and western groups following multiple vicariance events that
shaped the current distribution pattern of Palaearctic lizards. High complexity
in the ecological niche evolution between the Mediterranean and non-Mediterranean
climatic regions was detected. The generally low niche divergence
among members of the eastern group and the remarkable climatic divergence
within the western group highlight an important role of temperature seasonality
in a Mediterranean and Atlantic climate context. The results also suggest
niche conservatism in terms of microhabitats described by vegetation cover.
Main conclusions The ocellated lizards provide an interesting example of a
vertebrate radiation where niche shift (with or without vicariance) and niche
conservatism alternate in different niche axes shaping current biogeographical
patterns.

Aim
Diversification and rapid radiation are well documented in lacertid lizards. Niche conservation is frequently observed among related taxa, whereby niches will not change much during speciation events. Here, we investigate the relationship between environmental niche divergence and phylogenetic relatedness in a widespread group of green lizards, the Lacerta trilineata group.
Location
Eastern Mediterranean, Anatolia and adjacent regions.
Methods
A dated phylogeny based on three mitochondrial genes was contextualized using species distribution models of all genetically identified lineages in the Lacerta trilineata group. Based on this analysis, ancestral climatic niche occupancy was reconstructed using niche occupancy profiles. Niche divergence among lineages was quantified by computing multivariate niche overlaps.
Results
All taxa are associated with humid areas, but there is extensive variation in their climatic niche breadths and positions, which accord with the main phylogenetic split in the group. Our results suggest divergent niche evolution within subclades and convergent evolution among clades, which implies only a limited degree of niche conservatism regarding annual variations in temperature and precipitation. In contrast, niche axes – mainly reflecting precipitation patterns of the coldest quarter – show a greater difference among clades than within clades, and therefore a higher degree of niche conservatism.
Main conclusions
Based on estimated divergence times between taxa and geological events in Anatolia, our results can be explained by fragmentation of the range of a hypothetical ancestral species, resulting in different adaptations of subclades either to humid continental climates or to more Mediterranean climates. Our study highlights deviations from classical niche conservatism theory due to significant niche shifts among sister taxa.

Diversification and rapid radiation are well documented in lacertid lizards. Niche conservatism is frequently observed among related taxa, whereby ecological niches remain mostly stable during speciation events. Here, we investigate the relationship between environmental niche divergence and phylogenetic relatedness in a widespread group of green lizards, the Lacerta trilineata group. A dated phylogeny based on three mitochondrial genes was contextualized using species distribution models of all genetically identified lineages in the Lacerta trilineata group. Based on this analysis, ancestral climatic niche occupancy was reconstructed using niche occupancy profiles. Niche divergence among lineages was quantified by computing multivariate niche overlaps. All taxa are associated with humid areas, but there is extensive variation in their climatic niche breadths and positions, which accord with the main phylogenetic split in the group. Our results suggest divergent niche evolution within subclades and convergent evolution among clades, which implies only a limited degree of niche conservatism regarding annual variations in temperature and precipitation. In contrast, niche axes – mainly reflecting precipitation patterns of the coldest quarter – show a greater difference among clades than within clades, and therefore a higher degree of niche conservatism. Based on estimated divergence times between taxa and geological events in Anatolia, our results can be explained by fragmentation of the range of a hypothetical ancestral species, resulting in different adaptations of subclades either to humid continental climates or to more Mediterranean climates. Our study highlights deviations from classical niche conservatism theory due to significant niche shifts among sister taxa.

While traditionally species recognition has been based solely on morphological differences either typological or quantitative, several newly developed methods can be used for a more objective and integrative approach on species delimitation. This may be especially relevant when dealing with cryptic species or species complexes, where high overallresemblance between species is coupled with comparatively high morphological variation within populations. Rock lizards, genus Darevskia, are such an example, as many of its members offer few diagnostic morphological features. Herein, we use a combination of genetic, morphological and ecological criteria to delimit cryptic species within two species complexes, D. chlorogaster and D. defilippii, both distributed in northern Iran. Our analyses are based on molecular information from two nuclear and two mitochondrial genes, morphological data (15 morphometric, 16 meristic and four categorical characters) and eleven newly calculated spatial environmental predictors. The phylogeny inferred for Darevskia confirmed monophyly of each species complex, with each of them comprising several highly divergent clades, especially when compared to other congeners. We identified seven candidate species within each complex, of which three and four species were supported by Bayesian species delimitation within D. chlorogaster and D. defilippii, respectively. Trained with genetically determined clades, Ecological Niche Modeling provided additional support for these cryptic species. Especially those within the D. defilippii-complex exhibit well-differentiated niches. Due to overall morphological resemblance, in a first approach PCA with mixed variables only showed the separation between the two complexes. However, MANCOVA and subsequent Discriminant Analysis performed separately for both complexes allowed for distinction of the species when sample size was large enough, namely within the D. chlorogaster-complex. In conclusion, the results support four new species, which are described herein.

While traditionally species recognition has been based solely on morphological differences either typological or quantitative, several newly developed methods can be used for a more objective and integrative approach on species delimitation. This may be especially relevant when dealing with cryptic species or species complexes, where high overall resemblance between species is coupled with comparatively high morphological variation within populations. Rock lizards, genus Darevskia, are such an example, as many of its members offer few diagnostic morphological features. Herein, we use a combination of genetic (two nuclear and two mitochondrial loci), morphological (15 morphometric, 16 meristic and four categorical characters) and ecological (eleven newly calculated spatial environmental predictors) criteria to delimit cryptic species within two species complexes, D. chlorogaster and D. defilippii, both distributed in northern Iran. Phylogenetic analyses of the molecular data confirmed the monophyly of D. chlorogaster, while D. defilippii is paraphyletic in respect to D. steineri. However, each of the complexes comprises several highly divergent clades, especially when compared to other congeners. We identified seven candidate species within each complex, of which three and four species are supported by Bayesian species delimitation within D. chlorogaster and D. defilippii (including D. steineri), respectively. Although the species within one complex lack clear diagnostic features, they can be well separated based on morphological variables when sample size is appropriate. Ecological Niche Modelling provided additional support for the identified species and niche overlaps between them are generally low, especially in the D. defilippii complex.

The oriental green lizards of the Lacerta trilineata group are widely distributed in Greece, Anatolia, the eastern Mediterranean, the southern Caucasus, and the Zagros mountains in Iran. We studied their phylogeography using three mitochondrial markers with comprehensive sampling from most representatives of the group. Their phylogeny and divergence times (implementing fossil-based molecular clock calibrations) were inferred using Bayesian methods, and haplotype networks were reconstructed to assess how genetic diversity and current distributional patterns were shaped. According to our phylogenetic analyses, the group constitutes a well-supported monophylum containing several distinct evolutionary lineages with high haplotype diversity. Vicariance might explain the divergences within most lineages that have accumulated by range restriction and expansion of populations as a result of Quaternary climate oscillations and glacial refugia. However, niche divergence appears to be a major force promoting speciation, and large scale distributional patterns between lineages were shaped earlier by multiple, independent dispersals out of Anatolia during the Pliocene and early Pleistocene. The results of the present study also suggest that the group is in need of a taxonomical revision because the identified lineages and genetic diversity are not congruent with the currently recognized subspecies.

Multiple dispersal out of Anatolia: biogeography and evolution of oriental green lizards. - In: Žagar, A. (ed.): Abstracts of the 8th International Symposium on the Lacertids of the Mediterranean Basin, 03 - 06 June 2013, Koper, Slovenia: 15-16.

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The oriental green lizards of the Lacerta trilineata group are widely distributed in Greece, Anatolia, the eastern Mediterranean, the southern Caucasus andthe Zagros mountains in Iran. We studied their phylogeography using three mitochondrial markers with comprehensive sampling from most representatives of the group. Their phylogeny and divergence times (implementing fossilbased molecular clock calibrations) were inferred using Bayesian methods, and haplotype networks were reconstructed to assess how genetic might explain the divergences within most lineages that have accumulated by range restriction and expansion of populations due to Quarternary climate oscillations and glacial refugia. However, niche divergence seems to be a major force promoting speciation and large scale distributional patterns between lineages were shaped earlier by multiple, independent dispersals out of Anatolia during the Pliocene and early Pleistocene. Our results also suggest that the group is in need of a taxonomical revision, as identified lineages and genetic diversity are not congruent with the currently recognised subspecies.

In this study a total of 29 Eremias strauchi strauchi specimens (10 males, 16 females and three juveniles), collected from different localities in northwest of Iran were examined. The study was based on morphological features including color pattern, morphometric measurements, pholidotic characters, as well as ecological and biological observations especially habitat, reproductive biology and distribution. Habitat features and new distribution localities were documented. Egg characters of Eremias strauchi strauchi such as shape, size, number and color are reported for the first time. Also, some other new information on this species has been given.

The Northwest of Iran, covering the three provinces of Ardabil, East Azerbaijan and West Azerbaijan, is a special area that, due to its particular geographical and climatic conditions has a rich biodiversity. The present study was conducted with the aim of studying the habitat, substrate type, feeding, conservation and distribution of lizards in this area from 2003 to now. Our results have revealed that there are 20 lizard species belonging to 13 genera and 5 families in this area. Lacertid lizards have the highest species diversity with 12 species. We also found that high ranges of habitats with a different substrate types are inhabited by lizard species. The number of species in pine regions compared with other types of area is considerably greater. The conservation status of lizards is unknown and requires further study.

Ahmadzadeh, F. & Kiabi, B.H. & Kami, H.G. & Hojjati, V. (2008) -

A Preliminary Study of the Lizard Fauna and Their Habitats in Northwestern Iran. - Asiatic Herpetological Research, 11: 1-9.

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Northwestern Iran has unique geographical and climatic conditions that support a rich flora and fauna. In
view of the lack of in-depth studies on the lizards of the region, an investigation was started in the northern part of
Ardabil Province for an inventory of this component of the fauna and their habitats. Collections were made from
October 2003 to June 2005 and 165 specimens were collected and identified. Five families, 12 genera and 15 species
are represented, including Agamidae: Laudakia caucasia, Phrynocephalus persicus, Trapelus ruderatus; Lacertidae:
Lacerta media media, Lacerta strigata, Lacerta brandtii, Darevskia raddei raddei, Eremias strauchi strauchi,
Eremias arguta, Ophisops elegans; Scincidae: Mabuya aurata transcaucasica, Eumeces schneiderii princeps,
Abelepharus bivittatus; Anguidae: Pseudopus apodus and Gekkonidae: Cyrtopodion caspium caspium. Comparing
this list to the data provided by Anderson (1999), it seems that most of the lizards are being reported for the Province
for the first time. The families Gekkonidae and Anguidae are newly recorded, and the gecko Cyrtopodion caspium is
first recorded from the west and northwest of Iran. With seven species represented in the area, lacertids have the highest
species diversity among the lizard families and need further study. Habitat features also have been given for all
species.

The evolutionary history of two lizards (Squamata: Lacertidae) is linked to the geological development of Iran. - Zoologischer Anzeiger, Leipzig, 270: 49-56.

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The Iranian Plateau is an area of highly complex landscape and incredibly diverse habitats, from salt deserts to temperate forests. Such a heterogeneous environment, at the crossroads of Asia, Europe, and Africa, fuels the high diversity and uniqueness of the Iranian fauna and flora. Nevertheless, our understanding of the historical processes that shaped the Iranian biodiversity remains limited. In this context, we revisit the evolutionary history of the two most widespread lizard species on the Iranian Plateau (Eremias persica and Mesalina watsonana) to explore whether these two taxa of similar ecology have been subjected to similar historical processes. To achieve that, we performed a series of phylogenetic analyses and thoroughly tested all possible alternative topologies based on available mitochondrial sequences (cytochrome b and 12S ribosomal RNA). Additionally, we estimated the number of mitochondrial clusters based on a novel single-locus delimitation method and the time of their divergence, using recently inferred evolutionary rates for lacertid species. The results indicate that the two taxa have been shaped by similar physiographic and climatic barriers since both phytogenies split into similar geographical clusters. However, E. persica has twice as long evolutionary time on the Iranian Plateau than M. watsonana. The first divergence of E. persica was estimated in the Middle Miocene (~13 Mya), at the time of the major uplift of the Zagros mountain range. The Upper Miocene tectonic rearrangement that resulted in the uplift of the Alborz mountains coincides with the original split in M. watsonana (~6.6 Mya) and several contemporaneous splits within E. persica.

Study on the Karyotype and C-banding pattern of chromosomes of Eremias argus argus. - Journal of Xuzhou Normal University, 1992-02.

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E · a · argus possesses 2u = 38 chromomes with a chromosome formula as 2n = 341 + 2m+ZW.NF=38. The C-bands of centric regions are found on all of chromosomes. Telomeric C-bands can be seen of pairs -1 chromosomes. As compared with species of close relative Eremias brenchleyi .there is a world of difference between the two state chearly :The Karyotype of E · a · argus more evolution. Meanwhile conduct a discussion on the relation of Karyotype evolution and species.

The urbanization process causes deep changes in the functioning of ecosystems, and has a negative effect on urban-wildlife’s health. Moreover, cities represent an important point of entry for exotic species, which can originate ecological problems. Reptile and amphibian populations have decreased in the last decades, and they can be used as environmental index because of their distribution along urban-natural gradients and their limited dispersal ability. In this study, an inventory of reptiles and amphibians was made in the city of Bilbao and later urban impact was determined by measuring the health status (immune response and parasite load) in 4 wall-lizard (Podarcis muralis) populations captured along an urban-natural gradient. Results showed a higher number of exotic herp species compared to native species in the city. We found that immune response increased as urbanization degree decreased. Finally, parasite load was higher in individuals from natural-deteriorated population.

We studied the female reproductive cycle in a population of the oviparous lizard Acanthodactylus schreiberi syriacus from sandy beaches of Tyre, Lebanon. Females emerged from hibernation in mid-April, one month later than males. They reached sexual maturity at 57 mm snout-vent length and had smaller body sizes than males. Fertilization started in May. Among 33 gravid females, only one female showed evidence for two clutches produced in the same reproductive season whereas most of females laid one clutch per season. Mean clutch size was 2.5 ± 1.0, ranging from 1-4. We found no significant correlation between female body size and clutch size.

The feeding biology of Ophisops elegans inhabiting Bursa was investigated. Stomach contents of 66 adult (28 ðð, 38 ¦¦)
specimens collected from 3 localities were investigated by numerical analysis. Most of the foods of the Ophisops elegans specimens
were insects (64.58%), the majority of which were of larval form (19.44%). Two specimens that were collected from Grsu were
maintained in a terrarium to observe their feeding behaviors.
During observations, Ophisops elegans exhibited various feeding behaviors depending on the type of prey and sometimes bit and ate
pieces of its prey (legs, wings, etc.).

Endogenous stages of Isospora acanthodactyli from sandy fringed-toed lizard (Acanthodactylus schmidti) were described for the first time, in Al-Qassim, Saudi Arabia. Merogony and gamogony took place inside the host cell’s cytoplasm, in the distal part of the small intestine. Multinucleated meronts, microgamonts and macrogametes were described and measured.

Al-Badry, K.S. (1975) -

Haematological studies on some reptiles from Kuwait. Part 3. Some corpuscular constants, blood glucose, total plasma protein and electrophoretic examination of blood proteins of the lizards Acanthodactylus scutellatus and Eremias brevirostris. - Kuwait Journal of Science, 2 (1): 159-165.

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Corpuscular Constants: hemoglobin, hematocrit (packed cell volume), and red cell count; also blood glucose and total plasma protein of the lizards Acanthodactylus scutellatus and Eremias brevirostris were determined. For Acanthodactylus scutellatus the mean values are : 8.5 g/100 ml, 30.0%, 1.135 x 106/mm3, 224.6 mg/100 ml, and 4.7 g %. For Eremias brevirostris the mean values are: 9.2 g/100 ml, 30.8%, and 1.424 x 106/mm3, 217 mg/100 ml, and 4.0 g % respectively. Electrophoretic mobilities of the three protein systems, serum, plasma and hemoglobin ere similar in the two lacertid lizards, but different from those previously obtained by the same author for two agamid lizards. Electrophoretic patterns of serum and plasma proteins consist in both lizards of four fractions, albumin, ل-globulin, â-globulin and م-globulin. The ل-globulin fraction was not resolved into and ل1 and ل-2- C12 as obtained in the agamid lizards. Regarding the relative proportions of the different protein fractions and the albumin: globulin ratio. both lizards showed but little variation from each other. In all individuals of the two species examined, no fibrinogen fraction was detected in their plasma patterns. Hemoglobin of both lizards behaved as a homogeneous single fraction slightly moving towards the anode. The rate of movement was nearly the same in both lizards, but it differed from that recorded for the agamid lizards.

Al-Badry, K.S. (1978) -

Activity and heath death of the lizard Acanthodactylus scutellatus with reference to the influence of the parietal eye. - Proceedings of the Egyptian Academy of Sciences, 28: 139-144.

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Al-Bakry, A.M. & Abdeen, A.M. & Abo-Eleneen, R.E. (2013) -

Comparative study of the osteology and locomotion of some reptilian species. - International Journal Of Biology and Biological Sciences 2 (3): 040-058.

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The aim of this study is to show the osteological characters of the fore- and hind-limbs and the locomotion features in some reptilian species: Laudakia stellio, Hemidactylus turcicus, Acanthodatylus scutellatus, Chalcides ocellatus, Chamaeleo chamaeleon, collected from different localities from Egypt desert and Varanus griseus from lake Nassir in Egypt. In the studied species, the fore- and hind-feet show wide range of variations and modifications as they play very important roles in the process of jumping, climbing and digging which suit their habitats and their mode of life. The skeletal elements of the hand and foot exhibit several features reflecting the specialized methods of locomotion, and are related to the remarkable adaptations. Locomotion is a fundamental skill for animals. The animals of the present studies can take various forms including swimming, walking as well as some more idiosyncratic gaits such as hopping and burrowing.

Al-Barazengy, A.N. & Salman, A.O. & Abdul Hameed, F.T. (2015) -

Updated list of amphibians and reptiles in Iraq 2014. - Bulletin of the Iraq natural History Museum, 13 (4): 29-40.

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The present work provides a list of all amphibians and reptiles recorded from Iraq up to
2014. It includes 115 species (105 species of reptiles and 10 species of amphibians) dating back to 25 families (20 families of reptiles and 5 families of amphibians). Conservation status of each species was mentioned.

Al-Dokhi, O.A. (2006) -

Ultrastructure of Sperm Head Differentiation in the Lizard, Acanthodactylus boskinus (Squamata, Reptilia). - International Journal of Zoological Research, New York, 2: 60-72.

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The aim of the present study was to describe the sperm head differentiation in the lizardAcanthodactylus boskinus at the ultrastructural level. For this purpose, five adult male lizards were collected during April and May, 2004 from the desert at the north-east area of Riyadh city, Saudi Arabia. Testes of the lizards were processed for transmission electron microscopy. Thorough observation of the ultrastructural differentiation events of the sperm head has led to a concept of four sequential morphogenetic phases; Golgi phase, cap phase, nuclear elongation and chromatin condensation phase and maturation phase. The Golgi phase is initiated by the proliferation of Golgi complexes in the early spermatids to generate numerous microvesicles and the phase is terminated by the formation of a voluminous acrosomal vesicle. Flattening of the developed acrosomal vesicle over the anterior nuclear portion to form a cap associated with dissolution of the acrosomal granule are the features of the cap phase. Subsequent elongation of the spermatid nucleus accompanied with nuclear chromatin condensation are the features of the following phase. The final maturation phase is characterized by complete chromatin condensation and disappearance of the microtubule manchette. The general features of the differentiation process, which accord with the common criteria of other reptiles, as well as the existing differences are discussed.

The major ultrastructural features of the sperm tail differentiation in the lizard, Acanthodactylus boskinus (A. boskinus) have been described. The initial step is the caudal migration of the centrioles followed by insertion of the proximal centriole in the nuclear fossa and extension of the distal centriole to form the microtubular axoneme. Thereafter, tail differentiation involves the development of neck region and middle, main and end pieces. The later three tail pieces along their extension encompass the axonemal complex which reveals the typical (9+2) arrangement of microtubules. The axonemal complex in the middle piece is enveloped within two successive sheaths, the mitochondrial and fibrous ones while its extension in the main piece is only encircled by the fibrous sheath. The end piece only manifestes the existence of the axonemal complex surrounded by the plasmalemma of the fully differentiated spermatids.

Al-Hashem, A. (2009) -

Effects of oil pollution on the population size of the desert lizard Acanthodactylus scutellatus and their ant prey at Kuwait’s Al-Burgan oil field. - Journal of Biological Sciences, 9 (7): 682-686.

Desert ecosystems in Al-Burgan oil fields of Kuwait were contaminated by heavy metals and petroleum hydrocarbons due to oil spill generated by the Gulf War in 1990. Studying sand lizard (Acanthodactylus scutellatus) population and their ant prey in the years 2002 and 2003 to detect the effects of oil pollution is now a focus of study. Polluted sites with apparently different degrees of pollution (namely tar mat, soot and clear sites) were compared with control sites outside this region. Total lizard numbers were recorded by using transect method. Number of ants was recorded by walking the transects and counting ants present. The results showed no difference in lizard population between the different study sites in 2002 and 2003 by applying the transect method. No difference in ant populations between the different study sites in 2002 and 2003. Although, the mean estimated lizard numbers were lower at the tar mat sites, the ant number in this location was greatest, meaning that food availability was highest at these sites. This suggests any reduction in the numbers of lizards is unrelated to low resource availability. The lizard numbers at the tar mat sites could be depressed by some property of the pollutants.

Al-Hashem, M. & Brain, P. (2009) -

Effects of oil pollution on body size and weight of the sand lizard Acanthodactylus scutellatus at the greater Al-Burgan oil field in Kuwait. - Research Journal Environmental Toxicology, 3: 56-59.

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This study investigated the impact of oil pollution on morphological measurements in adult male and female sand lizards captured in locations with apparently different pollution levels. The results of this study confirmed that there is sexual dimorphism in body size, with males being generally larger than females at all the study sites. Adult male (but not female) lizards were generally bigger at the Tar mat and soot than the clear and control sites. The increase in body size and weight suggests that there is a greater availability of food for these somewhat territorial reptiles in both the Tar mat and soot sites. An alternative explanation is that the food resource is affected by oil pollution such that lizards consuming prey with high levels of fat accumulate more adipose tissue in their bodies.

Al-Hashem, M. & Brain, P.F. & Omar, S. (2007) -

Effects of oil pollution at Kuwait’s Greater Al-Burgan Oil Field on polycyclic aromatic hydrocarbon concentration in the tissues of the desert lizard Acanthodactylus scutellatus and their ant prey. - Ecotoxicology, 16: 551-555.

Oil pollution in desert locations in Greater Al-Burgan oil fields of Kuwait generated by
the 1990 Gulf war has changed field behaviour and morphology of the Fringe-toed Lizard Acan-
thodactylus scutellatus (Audouin, 1827). Lizards from the dark ‘tar mat’ locations are notably
darker in colour than those from the control sites. Consequently, A. scutellatus collected from ‘tar
mat’ and counterparts from lighter control sites were compared in laboratory studies where the
lizards could choose between a dark and a light side of an enclosure. The former lizards showed a
clear preference for darker substrates whereas the latter clearly preferred the light substrate. Dark
lizards on tar mat have more effective crypsis but also some advantages in terms of solar gain
(surprisingly, reptiles from tar mat sites are generally larger than counterparts from uncontami-
nated sites). It is uncertain whether the presence of dark substrates for more than a decade (this
study was conducted in 2003) has resulted in a selection of morphologically dark sand lizards
with a preference for dark surfaces or whether these reptiles simply use their skin chromatophores
to adapt to different surfaces but show a preference for substrates similar to their body colour-
ation.

An attempt was made to study the effects of oil pollution in a desert location (the Greater Al-Burgan oil fields, an area damaged in the second Gulf War) in Kuwait on the behaviour of the Sand lizard A, scutellatus. Polluted sites with apparently different degrees of contamination (namely tar mat, soot and clear sites) were compared with control areas outside this region. Between 2002 and 2003, ten lizards (5 of each sex) on each polluted and each control site were observed in the field at a time of the year when they were highly active. Air, substrate and burrow temperatures were recorded and lizards were monitored for their morning emergence times, as well as their basking and foraging activities. The present study confirmed that the morning emergence times and the basking behavior varied in sand lizards among the different pollution site categories. Physical changes in the tat mat sites caused the substrate temperatures in these locations to rise more quickly in the morning in response to solar gain than was the case in the other sites. This gives lizards in these locations the opportunity to emerge earlier and to start eating more quickly, giving them an energetic advantage (perhaps, in turn, influencing their rates of growth and fecundity). The cleat sites had the next earliest emergence and were the next hottest but it is difficult to account for this in terms of the physical characteristics of this site. The basking times were clearly shorter on the dark soot and tar mat sites that appeared to have higher solar gain than control or clear sites. There did not appear to be any obvious differences in foraging activity of lizards in the different locations. It appears that some aspects of simple behaviour in these lizards provides a reliable, noninvasive indices for assessing oil pollution in desert locations. The precise impact of these changes in these reptiles on their long-term viability needs to be evaluated.

Home range and vagility of the lizards Acanthodactylus schmidti and Acanthodactylus boskianus in Saudi Arabia. - Journal of Arid Environments, 16 (1): 79-86.

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Male lizards had larger home ranges than females and, in general, spring home ranges were greater than winter ones. Overlap in home ranges was not uncommon; extent of overlap varied greatly. Use of burrows by both species was similar but there was a difference in the use of habitats, eg A. schmidti were often seen basking in open areas, but A. boskianus were rarely seen on bare sand away from vegetation.

A total number of 70 herpetofaunistic species organised in 3 orders and 18 families were surveyed
during a period of two years (2005– 2007) in southern Jordan. The orders are (1) Ophidia has 7 families:
Leptotyphlopidae (1 species), Typhlopidae (2 species), Boidae (1 species), Colubdridae (17 species),
Atractaspididae (1 species), Elapidae (1 species) and Viperidae (5 species). (2) Sauria has 7 families: Gekkonidae
(12 species), Chamaeleonidae (1 species), Agamidae (8 species), Lacertidae (3 species), Sincidae (8 species),
Anguidae (1 species), Varanidae (1 species). (3) Testudines has 4 families: Cheloniidae (2 species),
Dermochetylidae (1 species), Emydidae (1 species) and Testudinidae (1 species). The species listed were all
resident and were mostly found throughout the year. The diversity of terrestrial and aquatic ecosystems in the
study area encouraged the occurrence of the species. The results reinforce the necessity of longterm inventory
plannings in order to understand the ecology and the dynamics of herpetofaunistic and other wildlife
communities in the study area. The over-increasing of human impact on the existing natural resources in the
southern Jordan has threatened the ecology diversity of wildlife species, where the populations of some
herpetofaunaistic species (especially frog species) and many reptilian species are declining in diversity status
and abundance. The author recommend at the end the improving cooperation of different parties to enhance
the public awareness and to implement environmental laws and legislation to conserve the sensitive and rare
species of herpetofauna components.

A total of 70 herpetofauna species from 3 Orders and 18 Families were surveyed during a period of two years (2005-2007) in Southern Jordan. The Ophidia included Leptotyphlopidae, Typhlopidae, Boidae, Colubdridae, Atractaspididae, Elapidae and Viperidae. The Sauria included Gekkonidae, Chamaeleonidae, Agamidae, Lacertidae, Sincidae, Anguidae and Varanidae. Testudines included Cheloniidae, Dermochelyidae, Emydidae and Testudinidae. The species listed were all resident and mostly found throughout the year. The diversity of terrestrial and aquatic ecosystems in the study area likely encouraged species presence. The results reinforce the necessity for long-term inventory planning in order to understand the ecology and the dynamics of herpetofauna and other wildlife communities in the study area. The increasing human impact on the existing natural resources in Southern Jordan has threatened the ecology and diversity of wildlife species to such a degree that populations of anurans and many reptiles are declining in diversity, status and abundance. The author recommends improving cooperation of different parties to enhance public awareness and to implement environmental laws and legislation to conserve sensitive and rare species of herpetofauna.

Al-Sadoon, M.K. (1987) -

The influence of temperature and activity on aerobic and anaerobic metabolism in the viviparous lizard, Lacerta vivipara Jacquin. - The Herpetological Journal, 1: 181-185.

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Aerobic and anaerobic metabolic rates were determined at temperatures between 20°C and 35°C for the viviparous lizard, Lacerta vivipara. Both parameters were found to be maximal around the preferred body temperature (30°C) with a low thermal temperature dependence above PBT. It is noted that L. vivipara does not need a large rate of anaerobic support and aerobic metabolism could supply the energy needed for activity.

The resting metabolic rate-temperature curves of juvenile, sub-adult and adult stages of the lizard Acanthodactylus schmidti were studied at 10, 15, 20, 25, 30 and 35°C. The mass-specific resting metabolic rate equations at different temperatures were calculated. Juveniles and sub-adults were found to have higher rates of O2 consumption than adults. The values for mass regression coefficient «b» proved to be temperature-dependent and ranged from 0.11 at 10°C to 0.58 at 35°C

Al-Sadoon, M.K. & Spellerberg, I.F. (1985) -

Effect of Temperature on the Oxygen Consumption of Lizards from different Climatic Regions. - Amphibia-Reptilia, 6 (3): 241-258.

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Oxygen consumption levels and metabolic rate temperature curves of various lizard species from three different climatic regions were examined in relation to ambient temperature. The species used in this research were as follows: Anguis fragilis, Lacerta vivipara, Lacerta agilis (cool temperate species); Blanus cinereus, Podarcis hispanica, Podarcis lilfordi brauni, Podarcis lilfordi lilfordi, Podarcis muralis, Psammodromus algirus, Tarentola mauritanica (warm temperate species); Chalcides ocellatus, Acanthodactylus opheodurus, Acanthodactylus schmidti (desert species). A double chamber volumetric closed system was used to measure the resting oxygen consumption of the lizards. Acute oxygen consumption determinations were made, that is the lizards were not allowed to acclimate to the test temperatures. Interspecific differences in levels of resting oxygen consumption and in the characteristics of the metabolic rate temperature curves were examined in relation to methods of thermoregulation and in relation to the ecology of the respective species. Evidence for `temperature dependent shifts` and `low thermal dependence` was found in the metabolic rate temperature curves of some species. A diminishing Q10 at or below the voluntary body temperatures suggests some degree of metabolic homeostatsis and energy conservation.

The ocellated lizard, Chalcides ocellatus, and the common lizard, Lacerta vivipara, were acclimated to two temperature regimes (10 and 30 degrees C with 12L:12D photoperiod) for 2 weeks. The oxygen consumption rates were measured for both species at temperatures between 5 and 35 degrees C. Cold acclimation was observed in L. vivipara and was in the form of an upward translation of the metabolic rate-temperature curve in comparison to the warm-acclimated lizards (30 degrees C). Chalcides ocellatus showed a response to cold acclimation by decreasing its metabolism only at 10 degrees C.

Isopora acanthodactyli, new species from Acanthodactylus schmidti with a new geographical record for I. deserti from Agama pallida (Finkelman and Paperna, 1994) in Saudi Arabia. - Pakistan Journal of Zoology, 29: 219-223.

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Isospora acanthodoclyli, new species is described from the feaces of Acanthodoctylus schmidti from Al-Thomama area in the central region of Saudi Arabia. Sporulated oocysts are spherical to subspherical 27.9×25.5 (25 1-29 8×22 7-27.8) μm Oocyst wall is smooth and bilayered Micropyle, oocyst residuum and polar granule absent Sporocysts are ovoid 14.6×11.1 (13.2-15.2×9 1-11.7) μm. Stieda and substiedal bodies are present. Sprozoites are hanana shaped with two spherical refractile globules. Isospora deserti (Finkelman and Paperna, 1994) is redescribed from the feces of Agama pallida in new geographical location. Sporulated oocysts measurement were similar to those reported previously from the same host in Israel (Finkelman and Paperna, 1994). Except for our observation of double layered wall, the oocyst morphology was identical to the original description of I. deserti.

An account of the Reptilia collected by Dr. F.P. Maynard, Captain A.H. McMahon, C.I.E., and the members of the Afghan-Baluch Boundary Comission of 1896. - Journal of the Asiatic Society of Bengal, 65: 550-566.

The composition of the food of Genetta genetta was studied from 17 groups of droppings with more than 2000 remains of animals and an undeterminated number of plants. On the small island of Cabrera lizards form the main diet. On the other islands mammals (Rodentia) and lizards (Lacertidae, Geckonidae) are the most common prey animals, followed by various insects. Plant material has been found more regularly on Ibiza. Genetta certainly is an euryphagous animal, but this behaviour pattern may alter to a stenophagous phase in certain environments.

Caves are prone to preserve fossils, mainly of vertebrates. On the Balearic Islands the Pliocene and Quaternary vertebrate fossil record allows to study the faunal composition and faunistical turnover on each island. On the Gymnesic Islands Myotragus balearicus was an endemic fossil Caprinae that lived until human arrival. It was the key species to understand the islands` ecology. The terrestrial vertebrate fauna that lived with Myotragus comprised a dormouse and a shrew, as well as an endemic lizard and the ferreret. The Upper Pleistocene and Holocene vertebrate fauna from the Pityusic Islands was only composed by birds, bats, and a lizard species. The absence of terrestrial mammals in this fauna constitutes the greatest faunistical anomaly in the whole Mediterranean.

In this paper we present a biogeographical analysis of the herpetological biota of the Balearics and Pityusics. Baleaphyne muletensis, Lacerta (Podarcis) lilfordi and Lacerta (Podarcis) pityusensis are considered as palaeoendemics and, plausibly, they colonized our islands during the Messinian. All the remaining species of the present day herpetological biota must have inmigrated to Balearics and Pityusiccs due to the intervention of man.

The habitat and systematics of Acanthodactylus schmidti Hass (1957) (Sauria, Lacertidae). - Journal of the College of Science. King Saud University, 19 (2): 89-95.

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This paper describes the habitat of Acanthodactylus schmidti Hass in Saudi Arabia and its present systematic position. The evidence suggests A. schmidti is a valid species. It is adapted to the sand dune habitat, particularly in areas with low scrubby vegetation. Its burrows are dug only among the roots of Hammada elegans .

Alekperov A.M. (1978) -

Amphibians and reptiles of Azerbaijan. - Elm, Baku. 264 pp.

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АЛЕКПЕРОВ, A.M. (1978) -

ЗЕМНОВОДНЫЕ И ПРЕСМЫКАЮЩИЕСЯ АЗЕРБАЙДЖАНА - ЭЛМ, БАКУ. 264 pp.

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Alekperov, A.M. (1971) -

On the distribution of the species Lacerta strigata on the Apsheron Peninsula. - Uchenye Zapiski Azerbaildzhanskogo Universiteta, Seriia biologicheskikh nauk, 1971 (3): 104-105.

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Alekperov, A.M. (1982) -

The present state of rare and threatened species and subspecies of herpetofauna in Azerbaidzhan. - Vertebrata Hungarica, Budapest, 21: 55-61.

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67 species and subspecies of amphibians and reptiles In the Azerbaldzhan Republic. 18 species and subspecies may be attributed the status of rare and disappearing (26.8%). Of this number, 8 species of amphibians and reptiles have been Included in the `Red Book of Rare Species of Animals and Plants of the USSR and those Endangered with Disappearance`: Pelobates syrlacus. Testudo graeca ibera, Agama ruderata, Lacerta parva, Elaphe longissima, Rhynchocalamus melanocephalus satunini, Vlpera ammodytes transcaucasiana and V. xanthina. The species are seriously endangered in the Azerbaldzhan: Triturus vulgaris, T. cristatus karelini, Bufo bufo verrucosissimus, Phrynocephalus heiloscopus, Mabuya aurata, Ablepharus bivittatus, Elaphe hohenackeri, Psammophis lineolatum, Vipera ursinii and Agkistrodon halys caucasicus.

We found significant variations in the pattern of sexual dimorphism in body size and shape among and within three lacertid species (Podarcis muralis, P. melisellensis and Archaeolacerta oxycephala), which live within a very restricted area (Lake Skadar, Montenegro) and under similar ecological settings. Each species has a specific pattern of size and shape relations between the sexes that proportionally changes with increase in size of the males, contrary to the expectations of Rensch`s rule. Our data do not confirm unequivocally that the level of size differences between the sexes (SSD) is affected by the body shape of two different morphotypes (cylindric vs. flattened body), or by the insularity conditions (island size, distance from mainland and the presence of syntopic lizard species). However, SSD appears to be positively correlated with clutch size at the interspecies level of comparisons.

We studied male and female one-year reproductive, fat body and liver cycle of the common wall lizard (Podarcis muralis muralis) from the urban and suburban localities of Belgrade. The minimum size at which females attain sexual maturity was between 49.78 and 51.25 mm of the snout-vent length. Vitellogenesis started at the end of March. The size and number of vitellogenic follicles varied considerably during the season. Increase in the female size leads to a significant increase in number but not in size of vitellogenic follicles. Females with oviductal eggs were found from late May to late July. Two clutches with mean size of 3.5±0.31 were laid in 1994. We found a significant positivecorrelationbetween female size and oviductalclutch size, while partial correlationbetween SVL and mean egg volume,as well as between clutch size and mean egg volume was insignificant. Relative oviductal clutch mass had the mean value of 0.17 ± 0.01, and was not correlated with female size. Testicles and epididymides exhibited maximal weight during April-June, and minimum in late July. The increase in male SVL lead to a significant increase in testicular and epididymal mass. Viable spermatozoa were produced from March were the smallest during the mating period while female fat bodies and liver decreased in weight during vitellogenesis. Mass of fat bodies and liver were significantly positively correlated with SVL in both sexes.

We estimated changes in the components of phenotypic variance and covariance among seven metric traits over the first 90 days of growth in the sharp-snouted rock lizard (Lacerta oxycephala). The broad-sense heritability estimated from the data representing mostly size components of the seven morphometic traits over four age periods was about 90%. The average broad-sense heritability for the shape components of the same traits and age periods amounted to about 42%. By using bivariate and multivariate statistical methods, ontogenetic allometry coefficients have been estimated. The average heritability estimated for the bivariate allometric coefficients was about 69%. Genetic correlation structures were highly integrated within each age period and this integration increased from newborn lizards to lizards aged up to 90 days. For absolute measurements, phenotypic correlation structures were less integrated than genetic correlation structures, while for size-free measurements the opposite trend was observed.

Alexander, G. & Marais, J. (2007) -

A Guide to the Reptiles of Southern Africa - Struik publishers, Cape Town.

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Alexander, G.J. & Horne, D. & Hanrahan, S.A. (2002) -

An evaluation of the effects of deltamethrin on two non-target lizard species in the Karoo, South Africa. - Journal of Arid Environments, 50 (1): 121-133.

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We measured the effects of deltamethrin, a pyrethroid insecticide, on individuals of two lizard species,Meroles suborbitalis and Pedioplanis namaquensis, in enclosures under controlled conditions, and under field conditions. We found significant effects on lizard mortality rates in the enclosure experiments, with most lizards dying within two months of treatment. Our field-based investigation, conducted over a five-month period, revealed significant reductions in abundance of M. suborbitalis (reduced by 52%) and P. namaquensis, (reduced by 72%) one and four weeks after spraying with deltamethrin. At 18 weeks after spraying there was no longer a significant difference in abundance measures between treatment and control plots, due to the invasion of the plots by juveniles.

Ontogeny of the conus papillaris of the lizard Gallotia galloti and cellular response following transection of the optic nerve: an immunohistochemical and ultrastructural study. - Cell and Tissue Research, 344 (1): 63-83.

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Spontaneous regrowth of the axons of retinal ganglion cells (RGC) occurs after unilateral optic nerve transection (ONT) in the lizard Gallotia galloti. We have performed an immunohistochemical and ultrastructural study of the conus papillaris (CP) of this lizard during ontogeny and after ONT in order to characterize its cell subpopulations, innervation and putative blood-brain barrier (BBB) and to evaluate changes occurring throughout regeneration. Proliferating PCNA+ cells were abundant between embryonic stage 33 (E33) and hatching. From E33, we observed Pax2+/GS+ glial cells in the primitive CP, which became increasingly pigmented and vascularised from E35. Conal astrocytes coexpressing Pax2 with vimentin and/or GFAP were identified from E37-E38. GluT-1+/LEA+/Pax2- endothelial cells (ECs) formed a continuous endothelium with tight junctions and luminal and abluminal microfolds. In adults, the peripheral blood vessels showed a thinner calibre, stronger GluT-1 staining and more abundant microfolds than those of the central CP indicating the higher specialization involved during transport within the former. Occasional pericytes, abundant Pax2+ pigment cells, LEA+ microglia/macrophages, unmyelinated Tuj1+ nerve fibres and SV2+ synaptic vesicles were also observed in the perivascular zone. After ONT, the expression of GluT-1 and p75NTR persisted in ECs, suggesting the preservation/early recovery of the BBB. Relevant ultrastructural alterations were observed at 0.5 months postlesion, although, by 3 months, the CP had recovered the ultrastructure of controls indicating tissue recovery. Abnormal newly formed blood vessels had developed in the CP-optic nerve junction. Thus, the CP is a central nervous system structure whose regenerating capacity might be key for the nutritional support of regenerating RGCs in G. galloti.

Identifying the factors generating ecomorphological diversity within species can provide a window into the nascent stages of ecological radiation. Sexual dimorphism is an obvious axis of intraspecific morphological diversity that could affect how environmental variation leads to ecological divergence among populations. In this paper we test for sex-specific responses in how environmental variation generates phenotypic diversity within species, using the generalist lizard Gallotia galloti on Tenerife (Canary Islands). We evaluate two hypotheses: the first proposes that different environments have different phenotypic optima, leading to shifts in the positions of populations in morphospace between environments; the second posits that the strength of trait-filtering differs between environments, predicting changes in the volume of morphospace occupied by populations in different environments. We found that intraspecific morphological diversity, provided it is adaptive, arises from both shifts in populations’ position in morphospace and differences in the strength of environmental filtering among environments, especially at high elevations. However, effects were found only in males; morphological diversity of females responded little to environmental variation. These results within G. galloti suggest natural selection is not the sole source of phenotypic diversity across environments, but rather that variation in the strength of, or response to, sexual selection may play an important role in generating morphological diversity in environmentally diverse settings. More generally, disparities in trait–environment relationships among males and females also suggest that ignoring sex differences in studies of trait dispersion and clustering may produce misleading inferences.

The present qualitative autoradiographic analysis aims to present the main features of morphogenesis and growth of claws in reptiles. Lizard embryos treated with tritiated thymidine reveal that epidermal cell proliferation in terminal digits is prevalent in the dorsal side and gives origin to the curved unguis of the claw. Less proliferation occurs in the ventral side of the digit tip where the concave sub-unguis is derived. Adult claws of a turtle show that thymidine-labelled cells are present along most of the epidermis of the claw, especially at the claw tip. Also, injection of tritiated histidine and proline, indicating active protein synthesis, confirm autoradiographic labelling along most of the epidermis of claws, in particular at the apical tip. The present study indicates that proximal matrix regions, as have been described in mammalian nails, are absent in reptiles. This pattern of claw growth probably derives from that of terminal digital scales. In fact reptilian (and avian) claws are formed from a modification of scales, a different condition from that present in mammals.

Alibardi, L. (2012) -

Observations on FGF immunoreactivity in the regenerating tail blastema, and in the limb and tail scars of lizard suggest that FGFs are required for regeneration. - Belgian Journal of Zoology, Brussels, 142 (1) : 23-38.

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Tail regeneration in lizards depends on the stimulation of growth factors, including Fibroblast
Growth Factors (FGFs). Light and ultrastructural immunolocalization of FGFs was compared between the regenerating
tail blastema and the limb where no regeneration occurs. A likely epithelial-mesenchymal transition
occurs following amputation in both tail and limb and FGFs are present in the wound epidermis of both organs
at 7-14 days post-amputation, and at lower intensity in mesenchymal cells of the blastema. Immunoreactivity for
FGFs disappears in the limb wound epidermis after 14 days post-amputation and in the epithelium covering tails
induced to form scars, whereas it remains in the apical tail epithelium. These observations suggest that scarring
in the limb or the induced scarring in the tail correlate with the disappearance of FGFs. Basic FGF is concentrated
in the incomplete basement membrane between the epidermis and the tail blastema where the essential
signaling process that allows the continuous growth of the regenerative blastema may occur. The study suggests
that the successful regeneration of lizard tail is dependent on the presence of FGFs in the wound epidermis,
which are probably released into the blastema.

In order to study some of the proteins involved in tissue remodeling and regeneration, the stem cell marker and dynamic cytoskeletal protein nestin have been detected using immunocytochemistry in normal and regenerating tail of lizard. In tissues of the normal tail, nestin is only present in the alpha-layer of scales and in sparse cells of connective tissues. However, during tail regeneration, nestin is more commonly detected in various tissues, especially in the epidermis and regenerating scales, muscles, nerves and in the growing spinal cord. Sporadic nestin immunopositive cells are seen in the regenerating blastema and derived connective tissues. In the epidermis, nestin is expressed in the main sites of cell proliferation located in the apical alpha-keratinocytes of the forming scales, but disappears in beta-cells. In the regenerating muscles, nestin is seen in myoblasts and elongating myotubes, but disappears in mature muscles. Nestin appears in the growing nerves and in a few neurons of the regenerating spinal cord. It is present in the apical ependymal cells and in peripheral nerves and Schwann cells. The dynamic cytoskeletal protein nestin appears to be activated in the main sites of cell proliferation and cytoskeletal remodeling needed for re-epithelization and the differentiation of new scales, muscles and nervous tissue.

Alibardi, L. (2014) -

Observations on Lumbar Spinal Cord Recovery after Lesion in Lizards Indicates Regeneration of a Cellular and Fibrous Bridge Reconnecting the Injured Cord. - Journal of Development Biology, 2( 4): 210-229.

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The lumbar spinal cords of lizards were transected, but after the initial paralysis most lizards recovered un-coordinated movements of hind limbs. At 25-45 days post-lesion about 50% of lizards were capable of walking with a limited coordination. Histological analysis showed that the spinal cord was transected and the ependyma of the central canal formed two enlargements to seal the proximal and distal ends of the severed spinal cord. Glial and few small neurons were formed while bridge axons crossed the gap between the proximal and the distal stumps of the transected spinal cord as was confirmed by retrograde tract-tracing technique. The bridging fibers likely derived from interneurons located in the central and dorsal grey matter of the proximal spinal cord stump suggesting they belong to the local central locomotory pattern generator circuit. The limited recovery of hind limb movements may derive from the regeneration or sprouting of short proprio-spinal axons joining the two stumps of the transected spinal cord. The present observations indicate that the study on spinal cord regeneration in lizards can give insights on the permissive conditions that favor nerve regeneration in amniotes.

Alibardi, L. (2015) -

Immunolocalization of a p53/p63-like preotein in the regenerating tail of the wall lizard (Podarcis muralis) suggests it is involved in the differentiation of the epidermis.. - Acta Zoologica, 2015. DOI 10.1111/azo.12130.

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During tail regeneration in lizards, the epidermis forms new scales comprising a hard beta-layer and a softer alpha-layer. Regenerated scales derive from a controlled folding process of the wound epidermis that gives rise to epidermal pegs where keratinocytes do not invade the dermis. Basal keratinocytes of pegs give rise to suprabasal cells that initially differentiate into a corneous wound epidermis and later in corneous layers of the regenerated scales. The immunodetection of a putative p53/63 protein in the regenerating tail of lizards shows that immunoreactivity is present in the nuclei of basal cells of the epidermis but becomes mainly cytoplasmic in suprabasal and in differentiating keratinocytes. Sparse labelled cells are present in the regenerating blastema, muscles, cartilage, ependyma and nerves of the growing tail. Ultrastructural observations on basal and suprabasal keratinocytes show that the labelling is mainly present in the euchromatin and nucleolus while labelling is more diffuse in the cytoplasm. These observations indicate that the nuclear protein in basal keratinocytes might control their proliferation avoiding an uncontrolled spreading into other tissues of the regenerating tail but that in suprabasal keratinocytes the protein moves from the nucleus to the cytoplasm, a process that might be associated to keratinocyte differentiation.

Alibardi, L. (2015) -

Regeneration of the epiphysis including the articular cartilage in the injured knees of the lizard Podarcis muralis. - Journal of Development Biology, 3 (2): 71-89.

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Cartilage regeneration is massive during tail regeneration in lizards but little is known about cartilage regeneration in other body regions of the skeleton. The recovery capability of injured epiphyses of femur and tibia of lizard knees has been studied by histology and 5BrdU immunohistochemistry in lizards kept at high environmental temperatures. Lizard epiphyses contain a secondary ossified center of variable extension surrounded peripherally by an articular cartilage and basally by columns of chondrocytes that form the mataphyseal or growth plate. After injury of the knee epiphyses, a broad degeneration of the articular cartilage during the first days post-injury is present. However a rapid regeneration of cartilaginous tissue is observed from 7 to 14 days post-injury and by 21 days post-lesions, a large part of the epiphyses are reformed by new cartilage. Labeling with 5BrdU indicates that the proliferating cells are derived from both the surface of the articular cartilage and from the metaphyseal plate, two chondrogenic regions that appear proliferating also in normal, uninjured knees. Chondroblasts proliferate by interstitial multiplication forming isogenous groups with only a scant extracellular matrix that later increases. The high regenerative power of lizard articular cartilage appears related to the permanence of growing cartilaginous centers in the epiphyses of long bones such as those of the knee during adulthood. It is likely that these regions contain resident stem cells that give rise to new chondroblasts of the articular and metaphyseal cartilage during most of the lizard’s lifetime, but can produce an excess of cartilaginous tissues when stimulated by the lesion.

After lumbar spinal cord transection, lizards recover some un-coordinated movements of the hind limbs including some walking ability. The transected spinal cord was examined using electron microscopy to study the degree of regeneration. A bridge tissue located between the two proximal and distal ends of the transected spinal cord, representing the regenerated cord, was found at 29-45 days post-lesion. The ependyma formed ampullar enlargements but no continuous central canal between the two stumps was re-established. Some cerebro-spinal fluid contacting neurons and peri-ependymal small neurons contacted by few synaptic boutons were found. Pale cells of astrocyte type were also identified. Numerous unmyelinated and sparse myelinated axons in the bridge appear connected to ependymal cells and oligodendrocytes. The origin of these axons remains unknown but previous studies have indicated that at least some of them are derived from interneurons, including those located around the central canal and in the dorsal grey matter of the proximal spinal cord stump. This limited regeneration can explain the recovery of part of the motor activity of the hind limbs in these lizards, possibly through the re-connection across the bridge of the intrinsic circuit of the central locomotory pattern generator.

The epiphysis of femur and tibia in the lizard Podarcis muralis can extensively regenerate after injury. The process involves the articular cartilage and metaphyseal (growth) plate after damage. The secondary ossification center present between the articular cartilage and the growth plate is replaced by cartilaginous epiphyses after about one month of regeneration at high temperature. The present study analyzes the origin of the chondrogenic cells from putative stem cells located in the growing centers of the epiphyses. The study is carried out using immunocytochemistry for the detection of 5BrdU-labeled long retaining cells and for the localization of telomerase, an enzyme that indicates stemness. The observations show that putative stem cells retaining 5BrdU and positive for telomerase are present in the superficial articular cartilage and metaphyseal growth plate located in the epiphyses. This observation suggests that these areas represent stem cell niches lasting for most of the lifetime of lizards. In healthy long bones of adult lizards, the addition of new chondrocytes from the stem cells population in the articular cartilage and the metaphyseal growth plate likely allows for slow, continuous longitudinal growth. When the knee is injured in the adult lizard, new populations of chondrocytes actively producing chondroitin sulfate proteoglycan are derived from these stem cells to allow for the formation of completely new cartilaginous epiphyses, possibly anticipating the re-formation of secondary centers in later stages. The study suggests that in this lizard species, the regenerative ability of the epiphyses is a pre-adaptation to the regeneration of the articular cartilage.

Alibardi, L. (2017) -

Immunohistochemical and western blot analysis suggest that the soluble forms of FGF1-2 and FGFR1-2 sustain tail regeneration in the lizard. - Annals of Anatomy – Anatoiscjher Anzeiger, 214: 67-74.

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Fibroblast Growth Factors 1-2 (FGF1-2) stimulate tail regeneration in lizards and therefore the distribution of their receptors, FGFR1-2, in the regenerating tail of the lizard. Podarcis muralis has been studied using immunofluorescence and western blotting. Immunoreactive protein bands at 15–16 kDa for FGF1-2 in addition to those at 50–65 kDa are detected in the regenerating epidermis, but weak bands at 35, 45 and 50 kDa appear from the regenerating connective tissues. Strongly immunolabeled bands for FGFR1 at 32, 60, and 80 kDa and less intense for FGFR2 only appear in the regenerating tail. In normal tail epidermis and dermis, higher MW forms are present at 80 and 115–140 kDa, respectively, but they disappear in the regenerating epidermis and dermis where low MW forms of FGFR1-2 are found at 50–70 kDa. Immunolocalization confirms that most FGFR1-2 are present in the wound epidermis, Apical Epidermal Peg, ependymal tube while immunolabeling lowers in regenerating muscles, blastema cells, cartilage and connectives tissues. The likely release of FGFs from the Apical Epidermal Peg and ependyma and the presence of their receptors in these tissues may determine the autocrine stimulation of proliferation and a paracrine stimulation of the blastema cells through their FGF Receptors.

Tissue regeneration in lizards represents a unique model of regeneration and scarring in amniotes. The tail and limb contain putative stem cells but also dedifferentiating cells contribute to regen- eration. Following tail amputation, inflammation is low and cell proliferation high, leading to regeneration while the intense inflammation in the limb leads to low proliferation and scarring. FGFs stimulate tail and limb regeneration and are present in the wound epidermis and blastema while they disappear in the limb wound epidermis 2–3 weeks postamputation in the scarring outgrowth. FGFs localize in the tail blastema and the apical epidermal peg (AEP), an epidermal microregion that allows tail growth but is absent in the limb. Inflammatory cells invade the limb blastema and wound epidermis, impeding the formation of an AEP. An embryonic program of growth is activated in the tail, dominated by Wnt-positive and -negative regulators of cell prolif- eration and noncoding RNAs, that represent the key regenerative genes. The balanced actions of these regulators likely impede the formation of a tumor in the tail tip. Genes for FACIT and fib- rillar collagens, protease inhibitors, and embryonic keratins are upregulated in the regenerating tail blastema. A strong downregulation of genes for both B and T-lymphocyte activation suggests the regenerating tail blastema is a temporal immune-tolerated organ, whereas a scarring pro- gram is activated in the limb. Wnt inhibitors, pro-inflammatory genes, negative regulators of cell proliferation, downregulation of myogenic genes, proteases, and oxidases favoring scarring are upregulated. The evolution of an efficient immune system may be the main limiting barrier for organ regeneration in amniotes, and the poor regeneration of mammals and birds is associated with the efficiency of their mature immune system. This does not tolerate embryonic antigens formed in reprogrammed embryonic cells (as for neoplastic cells) that are consequently elimi- natedimpedingtheregenerationoflostorgans.

Alibardi, L. (2018) -

Immunodetection of High Mobility Group Proteins in the regenerating tail of lizard mainly indicates activation for cell proliferation. - Acta Zoologica, 2018. https://doi.org/10.1111/azo.12259

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Immunodetection of High Mobility Group Proteins (HMGs) in the regenerating tail of lizard indicates activation for cell proliferation. Acta Zoologica (Stockolm). High Mobility Group Proteins (HMGs) are involved in chromatin assembling and control of transcription, especially during development. Transcriptome data indicate that HMGs are abundantly expressed in the early regenerating tail of lizards but their cellular localization remains unknown. Protein bands at 60–62 and 28–30 kDa are detected in western blots, more intense in the regenerating blastema. Immunodetection of HMGs in regenerating tail of the lizard Podarcis muralis indicates that these proteins are mainly localized in tissues where cell proliferation is as active as the apical wound epidermis, ependyma of the spinal cord and pro‐muscle aggregates. Few immunolabelled cells were seen in the regenerating cartilage and growing myomeres or nerves. Only in the wound epidermis labelled cells show a prevalent nuclear labelling while in other tissues a cytoplasmic labelling is prevalent. Only sparse immunolabelled cells are observed in the apical mesenchymal blastema and in the derived connective tissues formed in the mature regions of the regenerating tail, also destined to form adipose cells. The study suggests that the tissues with the highest immunolabelling correspond to those containing more proliferating cells, and that HMGs are mainly activated in these cells to promote cell division for the growth of the new tail.

The distribution of three anti-cytokeratin (-keratin) antibodies (AE1, AE2, AE3) in the epidermis of a lizard has been studied by immunocytochemistry at light and electron microscope and by immunoblot analysis. This study shows the expression of different keratins in the resting stage epidermis of the lizard Podarcis sicula. In this stage the epidermis has an external -layer, an underlying -layer, some layers of living suprabasal cells and a basal stratum germinativum. The AE1 antibody is localized in the basal and suprabasal cells only in the outer scale surface, but is absent from the inner surface, the hinge region and from the keratinized - and -layers. The AE2 antibody is mainly localized at the level of the hinge region and of the -layer and gives a lower reaction in the -layer. The AE3 antibody is mainly localized in basal and suprabasal cells, lower in the -layer, and absent from the -layer. The electron microscope shows that all the three antibodies immunolabel cytoplasmic fibrillar structures in the deep -layers and that AE2 and AE3 antibodies label small electron-dense areas in the external dense -layer within the electron-lucid matrix. Immunoblot analysis of the keratins extracted and separated by gel electrophoresis demonstrates the presence of a band of high molecular weight (67-68 kDa) positive to all three antibodies. In addition AE1 antibody recognizes a 44-45 kDa band and a 57-58 kDa band, AE2 recognizes a 60-61 kDa band, and AE3 recognizes a 47 kDa and a 56-57 kDa band. The localization of the keratins identified by immunoblot analysis in the epithelial layers is discussed taking in account the immunolabeling at light and electron microscope. The present study suggests that also in the normal epidermis of this reptiles, in both the - and the -layer, the molecular masses of keratins increase from the basal to the keratinized layers, a phenomenon which is generalized to adult and embryonic amniotes epidermis.

Using immunocytochemistry at light- and electron-microscope levels, we studied the distribution of three monoclonal antibodies (AE1, AE2, AE3) specific for mammalian α-keratins in regenerating lizard epidermis. We also characterized the keratins expressed during this process by immunoblotting after electrophoretic separation. The AE1 antibody is localized in the basal and suprabasal layers of prescaling and scaling epidermis. During the first stages of scale neogenesis, the AE1 antibody also marks the differentiating oberhautchen and β-layer, but it disappears from these layers as they mature. This antibody does not stain the prekeratinized and keratinized outermost layers in the hinge region. The AE2 antibody labels the superficial wound epidermis, prekeratinizing and keratinized β- and α-layers, but not basal and suprabasal cells. The AE3 antibody labels all living and keratinized epidermal layers, although AE3 immunoreactivity decreases and disappears as the β-layer matures. The ultrastructural study shows that the AE2 and AE3, but not the AE1, antibodies specifically label small electron-dense areas within the β-layer, suggesting retention of α-keratins. In the stages of tail regeneration examined, immunoblotting with the three antibodies used for the immunolocalization gives a pattern similar to that of the normal epidermis, except distally, where the process of scale differentiation begins. In this region, in addition to the keratin forms discovered in the normal and in proximal regenerating epidermis, an intense low molecular weight band at 40–41 kDa, positive to all three antibodies, is clearly detectable. Furthermore, in the distal region AE1 and AE3 antibodies, but not the AE2, recognize a weak band at 77–78 kDa not present in the normal and proximal epidermis. The localization and the possible role of the different keratins in the regenerating epidermis is discussed.

During the regeneration of lizard tail, nerves sprouting from ganglia and the spinal cord invade the blastema as far as the apical epidermis. Electron microscopical observations reveal axons storing dense granules (dg) and dense core vesicles (dcv) which are concentrated in nerve terminals or in axoplasmatic regions. In the regenerating spinal cord (SC) these terminals resemble aminergic-peptidergic endings and grow as far as the distal portion of the SC, which is made up of irregularly arranged ependymal cells. Some axons storing dcv contact blastematic cells and other nerve terminals show a plasma membrane incomplete or broken. Whether this latter aspect is due to fixation artifacts or physiological rupture is unknown. Nerves containing dcv and a few dg also originate from spinal ganglia innervating the regenerating tail. The accumulation of material into these endings is probably slow and a possible trophic influence on the regeneration of lizard tail is discussed.

In the present study, it was hypothesized that micromorphology of the surface of many lizard scales appears to mimic the topography of the habitat in which they live. Many authors have suggested that the microstructure of the superficial surface of scales have undergone important adaptations and have functional value in lizards. In this study, we investigated the variation and adaptation of the micromorphology and microstructure of the superficial surface of the dorsal and ventral scales from the mid-body region of Stellagama stellio (Agamidae), Stenodactylus petrii (Gekkonidae), Acanthodactylus boskianus (Lacertidae), Eumeces schneideri (Scincidae), Trachylepis quinquetaeniata (Scincidae), Scincus scincus (Scincidae), Varanus griseus (Varanidae), Chameleo chameleon (Chamaeleonidae). Skin specimens were prepared and analyzed using scanning electron microscopy. The dorsal and ventral scale surfaces had microstructure in the studied species and they exhibited unique patterns that somewhat resembled the topography of the microhabitats in which they lived. Similarity was detected in the three most related species, those having a common family, Scincidae. Ecomorphological relationships were detected between the dorsal and ventral scale microstructures and microhabitats. We conclude that environmental factors have observable influences on the microstructure of lizard scales.

Allan, G.M. & Prelypchan, C.J. & Gregory, P.T. (2006) -

Population profile of an introduced species, the common wall lizard (Podarcis muralis), on Vancouver Island, Canada. - Canadian Journal of Zoology, 84 (1): 51-57.

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Introduced species represent one of the greatest potential threats to persistence of native species. Therefore, it is important to understand the ecology of introduced species in order to develop appropriate mitigation strategies if required. In this study, using data collected in 1992-1993, we describe some fundamental population attributes of common wall lizards, Podarcis muralis (Laurenti, 1768), of Italian origin, introduced near Victoria, British Columbia, in the early 1970s. Male and female wall lizards reached similar snout-vent lengths, but males had relatively longer tails and were heavier. However, when gravid, females attained a body mass similar to that of males of equal snout-vent length. We found gravid females in all months from May to July, inclusive, but hatchlings did not appear in the field before late July. Growth rate was inversely related to body size, and lizards probably reached maturity in their second full summer. Larger lizards were moiré likely than smaller lizards to have experienced tail loss prior to capture, but the probability of tail loss upon capture was higher for smaller lizards than for adults. Our results suggest no fundamental differences in population characteristics between P. muralis on southern Vancouver Island and population at sites within the species natural range in Europe. Whether P. muralis on Vancouver Island is a threat to the native northern alligator lizard, Elgaria coerulea (Wiegmann, 1828), remains an open question.

Allan, M. & Prelypchan, C.J. & Gregory, P.T. & Waye, H.L. (1993) -

The ecology of introduced Common Wall Lizards (Podarcis muralis) on Vancouver Island, British Columbia. - Report submitted to the British Columbia Ministry of the Environment.

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Allen, A. (1975) -

Changes in population density of the Eyed lizard, Lacerta lepida, at three localities in Portugal between 1969 and 1975. - British Journal of Herpetology, 5: 661-662.

Comparative Phylogeography and species delimitation of the Arabian Peninsula lizards. - A thesis submitted to Bangor University for the degree of Doctor of Philosophy. Bangor University School of Biological Sciences. 167 pp.

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The Arabian Peninsula is an interesting area for research. This area has a complex geological history, a range of habitats and a number of known endemic species. However, to date, this area and its associated biodiversity has been poorly studied. The aims of this study were to investigate the phylogeography and to determine the species delimitation of a selected number of lizard species from the Arabian Peninsula. The phylogeography of fourteen co-distributed lizard species occurring within the Arabian Peninsula was investigated using a multispecies tree in STAR BEAST (*BEAST) to determine the divergence times and spatial patterns of the co-distributed species. Several common spatial and temporal patterns were identified among the different Arabian Peninsula species. The common patterns indicated close phylogeographic relationships between different regions and species. Importantly, these common patterns also corresponded to historical biogeographic processes. A wide range of ecological habitats was also detected for these groups of lizards and this was assumed to play a major role in establishing the current diversity and distribution patterns. In addition to detecting common patterns, this study also provided valuable information about the unique phylogeographical patterns shown by some of the studied species. Finally, this study also revealed patterns that provided strong evidence for the presence of multiple cryptic species within a species complex. Species delimitation methods were subsequently applied to two species that had previously demonstrated the potential for cryptic species within Acanthodactylus boskianus and A. opheodurus. Using a combined approach of genetic distance, allele networks, and Bayesian Phylogenetic and Phylogeography (BPP) analysis, this study was able to identify candidate species within A. boskianus and A. opheodurus. The mitochondrial DNA tree revealed potential candidate clades within these two species. These candidate species clades were then further examined at two nuclear loci and congruence was observed between the two markers for these clades. This congruence between mitochondrial and nuclear loci strongly indicates the discovery of several new species within A. boskianus and A. opheodurus, however further research is needed to confirm this discovery. In conclusion, this study provides the most detailed insight - to date - on the phylogeography and species delimitation of Arabian Peninsula lizards and provides the most up to date assessment of the diversity of the lizards in this important region.

Soricid mammals and lizards are small-sized, insectivorous vertebrates that are widespread and abundant in Mediterranean habitats. Because of their broad sympatry and their diet similarity, these taxa have been suspected to compete for food. Therefore, co-occurrence patterns between these taxa were studied at 72 sites in southern Italy by means of trapping methods. The assemblages were quite distinct depending on the site bioclimate: for the Lacertidae, Podarcis siculus dominated in the thermo-Mediterranean sites and P. muralisin the temperate sites, whereas, for the soricids, Suncus etruscusand two species of Crocidurawere dominant in thermo-Mediterranean sites and three Sorexspecies in the temperate sites. The mean number of captured soricids was statistically higher in the temperate sites, and was positively related to the first component of a Principal Component Analysis summarizing three collinear study site variables (elevation, mean annual temperature, annual rainfall), the reverse being true for lizards. Co-occurrence analysis revealed that a non-segregated structure was present in the dataset, whereas a randomization algorithm showed that the assemblage of small mammals and lizards was non-randomly structured, with the frequency distribution of shrews being non-independent by site from that of lizards. However, when we divided the sites by their bioclimatic regime (thermo-Mediterranean versus temperate), the non-randomness of the community structure disappeared, thus demonstrating that interspecific competition was not the main force driving these assemblages of species. The number of shrews captured in each sampling site was however significantly negatively related to the number of lizards, this pattern being linked to the bioclimate of the various sampling sites. Overall, our data indicated that the assemblage of shrews and lizards was likely regulated essentially by local climate and not by synecological (interspecific competition) dynamics.

Poikilothermic animals show a strong dependence on environmental temperature. However, they are able to adjust their body temperature through behavioral and partial physiological thermoregulatory adjustments. In this way, they optimize the heat loss and gains to keep the body within a range of thermal preference and to allow maximizing their physiological efficiency. A group of Gallotia galloti lizards were placed in a terrarium with a wide thermal variation in different regions. In this way, the animals were able to select the most thermally comfortable region. However, they should leave this region to get food, which was placed in a cold region, with a temperature far away from the comfort temperature. The time-lapse thermographic cinematography recordings allowed recording the environmental and body temperatures as well as the variations in different parts of the animal body in function to the temperature of the environment. In this way, it was found that the cephalic temperature was maintained within the 33 – 37 range in spite of occasional visits, the coldest spot in the terrarium at 13ºC to get food. Most frequently, the animals preferred to stay in 25-30ºC regions, while maintaining a cephalic temperature of up to 10ºC higher.

Anatolacerta has included 3 species and one of them is
Anatololacerta anatolica (Werner 1900). This species has two
distinct subspecies respectively; Anatololacerta anatolica anatolica
and Anatololacerta anatolica aegaea (Eiselt and Schmidtler 1987).
The systematic of Anatololacerta based on morphology and which
is included species has not been studied karyotypically. Therefore
we aimed to report the karyotype of specimens and obtained
differences from 4 different populations of Anatololacerta
anatolica subspecies which is distributed in Western Anatolia. The
result showed that two subspecies possess 2N = 38 uniarmed
(subtelocentric) chromosomes that is 36 macrochromosomes and 2
microchromosomes.Also meiotic plates showed that 19 bilavents.

We surveyed the historical Faid Hema, Ha’il region in Saudi Arabia and identified 22 species of reptiles (17 lizards and five snakes). We recorded for the first time nine reptiles (six lizards and three snakes) in the Faid Hema. Of these, one lizard, Mesalina adramitana, and two snakes, Cerastes gasperetti and Walterinessia aegyptia, were reported for the first time for the entire Ha`il region. We recorded 50% of all species for the entire study area at one site, which may be because it had three different habitat types near it (the Jannin mountain, soft soil, and firm, sandy soil). The next highest number of species we found was at an open site with firm, sandy soils and with gravel (containing 45.5% of the total species recorded), followed by a site that contained only sand (36.4% of the total). The lowest number of species was reported at an agricultural site, with only 13.6% of the total species found.

There are three types of cells in the vomero-nasal organ of Lacerta sicula and Natrix natrix: receptor cells, supporting cells and basal cells. The receptor cells bear microvilli and no cilia. In Lacerta centrioles are lacking, indicating that the ciliary apparatus can have no essential significance in the transducer process. In Natrix centrioles occur in the deeper dendritic region. The structural constituents of the dendrites are mitochondria, microtubules and characteristic vesicles the properties of which are described. The perikarya which have uniform structure send off axons of about 0.2 μ diameter. The supporting cells show signs of a very moderate secretory activity, which is different among the species investigated. The microvilli of the supporting cells are not distinguishable from those of the receptor cells. The dendrites of the latter are completely isolated by the apical parts of the supporting cells. The sheet-like processes of the supporting cells contain strands of tonofilaments and do not cover the perikarya of the receptor cells completely. Thus adjacent sensory cells or dendrites and sensory cells are separated among themselves only by the normal intercellular space. The ratio of sensory cells to supporting cells is about 7∶1. The basal cells resemble the supporting cells and replace these in the lower portion of the epithelium. The typical cellular junctions between sensory cells and supporting cells are described. There are no true tight junctions in the vomero-nasal sensory epithelium, and they are most probably absent from the nasal mucosa too. This absence would seem to indicate special conditions for cellular communication and the accessibility of the intercellular space for certain molecules. There is no sign of regeneration of sensory cells. Both immature blastema cells and degenerating receptor cells are not discernible.

We studied, for the first time, the age structure of Eremias strauchi strauchi (Kessler, 1878) from a sample (18 adults) of a population living in the vicinity of Iğdır (eastern Turkey) using the skeletochronological method. The maximum observed longevity was 7 years for males and 5 years for females. The age of males ranged from 4 to 7 years (mean 4.91, n = 12), while that of females ranged from 4 to 5 years (mean 4.66, n = 6). The average snout–vent length (SVL) was 61.10 mm in males and 60.82 mm in females. The differences in mean age and SVL between sexes were not statistically significant. Intersexual differences in body size were male-biased, but this state was not statistically significant. Age and SVL were positively correlated in both sexes.

Altunisik, A. & Eksilmez, H. (2018) -

Demographic life history traits in a population of a critically endangered species, Darevskia dryada (Darevsky & Tuniyev,1997). - Animal Biology, 68 (1): 27-37.

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Knowing the age structure of endangered species is important in order to contribute to future conservation studies for such species. In this context, we investigated age structure, age at sexual maturity, potential reproductive lifespan and longevity in a population of the Charnali lizard, Darevskia dryada, an endangered species from Turkey. The results show that the Charnali lizard has a longer life span than other lizards of the genus Darevskia that live in the same region. We estimated that these lizards attain their sexual maturity at the age of one or two years and the potential reproductive life span for males and females is six and five years, respectively. As in many other lizards, the Charnali lizard exhibited a low-level male-biased sexual dimorphism in terms of increased size. We believe that this study, in which prior information related to the Charnali lizard is shared, will contribute to future conservation activities for this critically endangered species.

In this study, the age composition of a Podarcis tauricus population from Saros Bay (northwest Turkey) was determined by skeletochronology. 27 preserved adults (20 females ,5 males, 2 juveniles) were evaluated and the maximum observed lifespan was recorded to be 7 years in females and 6 years in males. The mean age was 5.2 ± 0.2 years in males and 5.0 ± 0.2 years in females; the mean snout-vent length (SVL) was 63.0 ± 0.7 mm (range: 60.4 – 64) in males and 61.3 ± 0.9 mm (range 52.4 – 68.5) in females. No significant differences were found between sexes in terms of body size, but the sexual dimorphism index (SDI), was determined as 0.03, indicating a weakly male-biased. Adult survival rate and adult life expectancy were estimated to be 0.51 and 2.54 for female individuals, respectively. The lizards reached sexual maturity between 2 and 3 years. No significant difference was found between the sexes in terms of age and head sizes. The age and SVL were correlated only in females. Additionally, a significant positive correlation was found among SVL, head length (HL), and width (HW) in females.

The life history characteristics of individuals may vary depending on the altitude gradient. In this study, some life history traits such as age structure, age at sexual maturity, lifespan, body size, head length and head width were tested for two different altitude populations of Ophisops elegans. The results suggest that the individuals from the high altitude (Sivas, 1595 m) population have a higher mean body size and longer and wider head structure than the low-altitude (Çanakkale, 7 m) population. Although, the mean age of the highland population was higher than the lowland population, this case was not found statistically significant. In this study, both females and males were estimated to reach sexual maturity at an average of 3 years. The expected positive correlation between age and body size was recorded for only females of the Çanakkale population.

Opposed elevational variation in prevalence and intensity of endoparasites and their vectors in a lizard. - Current Zoology, 2018, https://doi.org/10.1093/cz/zoy002

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Studying the causes of parasite geographic distribution is relevant to understand ecological and evolutionary processes that affect host populations as well as for species conservation. Temperature is one of the most important environmental variables affecting parasite distribution, as raising temperatures positively affect development, reproduction, and rate of transmission of both endo- and ectoparasites. In this context, it is generally accepted that, in mountains, parasite abundance decreases with elevation. However, empirical evidence on this topic is limited. In the present study, we analysed the elevational variation of haemoparasites and ectoparasites of a lizard, Psammodromus algirus, along a 2200-m elevational gradient in Sierra Nevada (SE Spain). As predicted, ectoparasite (mites, ticks, mosquitoes, and sandflies) abundance decreased with elevation. However, haemoparasite prevalence and intensity in the lizard augmented with altitude, showing a pattern contrary to their vectors (mites). We suggest that tolerance to haemoparasites may increase with elevation as a consequence of lizards at high altitudes taking advantage of increased body condition and food availability, and reduced oxidative stress. Moreover, lizards could have been selected for higher resistance against haemoparasites at lowlands (where higher rates of replication are expected), thus reducing haemoparasite prevalence and load. Our findings imply that, in a scenario of climate warming, populations of lizards at high elevation may face increased abundance of ectoparasites, with accompanied strong negative effects.

Icosahedral nucleo-cytoplasmic large DNA viruses (NCLDV)-like viruses, which form inclusions in the erythrocyte cytoplasm of reptiles, were previously presented as candidates for a new genus of the Iridoviridae family. The present work describes the distribution of infected lizard hosts and ultrastructural characteristics of the viral inclusions of NCLDV-like viruses from Portugal and adjacent locations in Spain. Giemsa-stained blood smears of 235 Lacerta schreiberi from Portugal and Spain, 571 Lacerta monticola from the mountain Serra da Estrela (Portugal), 794 Podarcis hispanica from several localities in Portugal and Spain, and 25 Lacerta dugesii from Madeira Island, were studied. Infection in L. schreiberi was only found in mountain populations, up to 30% in Serra da Estrela and 9–11% elsewhere. It was absent in lizards from lowlands. Prevalence of infection among L. monticola in Serra da Estrela was 10%; infected lizards were found during March to July and October but not in August and September. Infection in P. hispanica was below 3.3%. Only one infected specimen of L. dugesii was identified by light microscopy. Ultrastructural examination of infected samples revealed that the inclusions are virus assembly sites of icosahedral cytoplasmic iridovirus-like virions. Virions from different host species have different ultrastructural features and probably represent different related viruses.

Assessment of the Endangered Species Podarcis carbonelli on a Microgeographic Scale: A Molecular, Morphological and Physiological Approach. - Masters Thesis, Western Kentucky University

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The lizard Podarcis carbonelli is an endangered species endemic to the Iberian Peninsula. One location where this species occurs is at the Berlengas Natural Preserve, an Atlantic archipelago off the coast of Portugal. These island populations are geographically separated from nearby mainland populations. The fundamental question is, are these insular individuals distinct from the mainland populations? Four localities were chose for comparison: two island populations and two nearby coastal populations. We assessed this question using three distinct approaches: molecular, morphological and physiological approach. We sequenced the 12S RNA, the mtDNA Control Region and the 7th intron of the !-fibrinogen gene and determined genetic diversity values as well as several parameters of population structure and differentiation. Individuals from these populations were also measured for several biometric characters and their blood lactate concentration was sampled. There was no genetic variation in both the mtDNA regions analyzed. The nuclear intron revealed high levels of genetic variation, with islands having in general lower values than the mainland regions. The four populations sampled had low levels of divergence; the populations of Berlenga and Peniche were the most distinct and the populations of Farilhão and Baleal were the most similar from the four populations sampled. Morphometric analyses revealed a different pattern of similarity among populations with the population of Farilhão being the only population statistically distinct from all other populations based on mass and SVL. Furthermore, island populations were in general more similar to each other than to mainland populations, with the exception of Berlenga males which in size are more similar to the Peniche males. The analysis of the blood lactate concentration revealed that the population of Peniche has significantly lower blood lactate levels than the populations of Farilhão and Berlenga. The lack of genetic differentiation found in the populations under study is most likely due to the recent divergence of these populations. Furthermore, the genetically most different populations (Berlenga and Peniche) are not the most distinct in terms of morphology, particularly the males. This suggests that genetic drift, the most likely mechanism behind the genetic differentiation seen, is not responsible for the morphological differences observed. The morphological differences seen can be attributed to: a possible difference in age of the individuals in each population; mechanisms of natural selection that are favoring specific phenotypes in each of the populations, or phenotypic plasticity. The differences in blood lactate levels found between the population of Peniche and the island populations can be attributed to differences in predatory pressure or home range size. It is suggested that the island populations are closely monitored due to their likely isolation, low mtDNA diversity and possible higher predatory pressure than initially predicted.

The use of a lacertid lizard as a model for reptile ecotoxicology studies: Part 2 – Biomarkers of exposure and toxicity among pesticide exposed lizards. - Chemosphere, 87 (7): 765–774.

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As part of a wider study examining the impacts of corn pesticides on lacertid lizards in north-western Portugal, we examined various physiological, biochemical, and histological biomarkers of exposure and effect among field populations of Podarcis bocagei. Biomarkers included body condition index, standard metabolic rate, locomotor performance, parasitization, glutathione oxidative pathways and related enzyme activity, lipid peroxidation and liver and testis histology. Few of the various biomarkers investigated provided statistically significant evidence of toxic effect. However, using a weight of evidence approach, we conclude that pesticides are affecting lizards living in the vicinity of pesticide exposed corn agriculture sites. Lizards from these locations present a profile of animals under metabolic stress with reduced condition indices, increased standard metabolic rate, lower incidence of hepatocyte vacuolation, altered iron metabolism, increased activation of GSH oxidation pathways, and even increased prevalence of hemoparasites.

Mesocosms (i.e., outdoor, man-made representations of natural ecosystems) have seldom been used to study the impact of contaminants on terrestrial ecosystems. However, mesocosms can be a useful tool to provide a link between field and laboratory studies. We exposed juvenile lacertid lizards for a period of over one year to pesticides (herbicides and insecticides) in mesocosm enclosures with the intention of validating field observations obtained in a previous study that examined the effects of corn pesticides in Podarcis bocagei. Our treatments replicated field conditions and consisted of a control, an herbicides only treatment (alachlor, terbuthylazine, mesotrione and glyphosate) and an herbicides and insecticide treatment (including chlorpyrifos). We used a multi-biomarker approach that examined parameters at an individual and sub-individual level, including growth, locomotor performance, standard metabolic rate, biomarkers of oxidative stress, esterases and liver histopathologies. Although mortality over the course of the exposures was high (over 60%), surviving individuals prospered relatively well in the mesocosms and displayed a broad range of natural behaviours. The low numbers of replicate animals compromised many of the statistical comparisons, but in general, surviving lizards exposed to pesticides in mesocosm enclosures for over one year, thrived, and displayed few effects of pesticide exposure. Despite the difficulties, this work acts as an important stepping-stone for future ecotoxicology studies using lizards.

Mesocosms (i.e., outdoor, man-made representations of natural ecosystems) have seldom been used to study the impact of contaminants on terrestrial ecosystems. However, mesocosms can be a useful tool to provide a link between field and laboratory studies. We exposed juvenile lacertid lizards for a period of over one year to pesticides (herbicides and insecticides) in mesocosm enclosures with the intention of validating field observations obtained in a previous study that examined the effects of corn pesticides in Podarcis bocagei. Our treatments replicated field conditions and consisted of a control, an herbicides only treatment (alachlor, terbuthylazine, mesotrione and glyphosate) and an herbicides and insecticide treatment (including chlorpyrifos). We used a multi-biomarker approach that examined parameters at an individual and sub-individual level, including growth, locomotor performance, standard metabolic rate, biomarkers of oxidative stress, esterases and liver histopathologies. Although mortality over the course of the exposures was high (over 60%), surviving individuals prospered relatively well in the mesocosms and displayed a broad range of natural behaviours. The low numbers of replicate animals compromised many of the statistical comparisons, but in general, surviving lizards exposed to pesticides in mesocosm enclosures for over one year, thrived, and displayed few effects of pesticide exposure. Despite the difficulties, this work acts as an important stepping-stone for future ecotoxicology studies using lizards.

Lizards are one of the least studied groups in ecotoxicology and despite a recent increase in number of studies, there is still a lack of knowledge regarding their response to environmental contamination. Lacertid lizards have been identified as potential model species for reptile ecotoxicology in Europe. They offer several advantages over other groups as they are usually highly abundant; have strong site fidelity; are easy to capture, mark and recapture in the field; and are easy to maintain in captivity. The main goal of our project is to assess if highly abundant lacertid lizards belonging to the genus Podarcis in Southern Europe, can be used as bioindicators of pesticide exposure and toxicity in areas of intensive agriculture. We have chosen Podarcis bocagei because is a widely abundant species and the only lacertid present on high pesticide usage corn fields dominating the landscape of north- western Portugal. As a first approach, we will document the abundance, demography and biological fitness of the species occurring in areas of either intensive or negligible pesticide usage. A second approach aims to study naïve lizards in more controlled experiments, both in mesocosm and laboratory experiments to validate field observations. We intend to employ a variety of techniques to assess pesticide exposure and toxicity including biochemical, behavioural, morphological and molecular biomarkers. On a larger scale we hope to study bioaccumulation processes in lizards, reflect on the biological significance of the reported tissue concentrations and develop criteria to predict risk to lizard populations. In the future, similar projects using lacertids as bioindicators could be developed in other parts of the Mediterranean Basin.

The use of a lacertid lizard as a model for reptile ecotoxicology studies - Part 1 Field demographics and morphology. - Chemosphere, 87 (7): 757-764.

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At the European level, lacertid lizards have been proposed as potential model species for reptile ecotoxicology. We studied demographic and morphological aspects of natural field subpopulations of Podarcis bocagei inhabiting similar agricultural habitats which were either regularly exposed to pesticides, or not. Parameters examined in this study included population size and density, sex ratio, adult body size, fluctuating asymmetry in femoral pores and parasite prevalence. In general, we detected few statistically significant differences between the exposed and reference subpopulations. Although field situations are ecologically complex and factors other than pesticides may be acting, the absence of observable effects on field subpopulations is probably indicative that lizards are coping or compensating for this level of exposure.

In Europe, reptiles have been recently included in environmental risk-assessment processes for registration of plant-protection products. However, data on toxicity effects of most compounds are lacking. Chlorpyrifos is the most commonly used organophosphorus insecticide worldwide. In the present study, the authors exposed a lacertid lizard, Podarcis bocagei, to sublethal concentrations of chlorpyrifos. Individuals were exposed through spiked food for a period of 20 d (low dose 0.12 mg/kg/d, high dose 1.57 mg/kg/d). After exposure, various biomarkers of exposure and effect were evaluated, including the activities of glutathione S-transferase and enzymes involved in the glutathione redox cycle, glutathione concentrations, activities of esterases, liver and testes histopathologies, as well as locomotory and predatory behavior. The results indicate that sublethal, subchronic exposure to chlorpyrifos can affect P. bocagei in a dose-dependent manner. Adverse effects occurred at both the subindividual and individual levels, including inhibition of carboxylesterases and cholinesterases (ChEs), liver histopathological changes, and altered predatory behaviors. Animals exposed to chlorpyrifos took more time to capture and subdue prey items. The results suggest a link between effects at subindividual levels of organization with those observed at the whole individual level after exposure to environmentally realistic dosages of chlorpyrifos.

Lizards are among the least studied groups in ecotoxicology and despite a recent increase in the number of studies, there is still a lack of knowledge regarding their response to environmental contamination. In Europe, lacertid lizards have been identified as potential model species for reptile ecotoxicology. The main question of this project was to assess if a highly abundant lacertid lizard belonging to the genus Podarcis, could be used as a bioindicator of pesticide exposure and toxicity in agricultural areas. To achieve this end, we used a three-stage tiered approach. The first tier took the form of a field survey to document the nature of the exposure and the population parameters of lacertids occurring in areas of intensive pesticide usage as well as areas of negligible pesticide usage. The second tier consisted of a mesocosm study in which naïve lizards were exposed to pesticides in a controlled experiment. Finally, the third tier included a laboratory approach to the effects of one of the most common insecticides used worldwide, chlorpyrifos. We assessed pesticide impact in individuals from the different tiers using a comprehensive set of biomarkers applied at different levels of biological organization, including population parameters as well as behavioral, physiological, biochemical and histological biomarkers. We detected few statistically significant differences between reference and exposed populations in the field study. Confirming the difficulty to isolate the effects of contaminants on natural populations where other local factors, natural cycles or stochastic events occur. P. bocagei populations seem to be able to compensate with the observed level of pesticide exposure. Nevertheless, individuals living in exposed sites seem to be less ecologically fit than those living in reference sites, presenting a depleted nutritional status and signs of metabolic stress. The results from the mesocosm setting reinforce these results. Lizards prospered relatively well in all enclosures, treated or not with pesticides, presenting a broad range of natural behaviors. The laboratory approach confirmed P. bocagei as valuable indicator of sub-lethal exposure to environmental realistic doses of chlorpyrifos. According to our results, P. bocagei seems to be a suitable bioindicator of pesticide exposure.

Geographical variation in scalation traits of the Iberian green lizard Lacerta schreiberi was studied measuring 27 morphometric and pholidotic traits in 533 lizards from 12 populations. The results show a little morphological differentiation between the populations of the mountains of Central System and the Atlantic side of the Iberian peninsula. The reduced width of the occipital plate of Central System lizards appears as the most remarkable difference between this population and the other populations analysed. This differentiation is consistent with the presence of two groups of mitochondria DNA haplotypes in each of these geographic areas. Nevertheless, the differences detected were too insufficient to consider a modification of the taxonomic status of Lacerta schreiberi.

The persistence of populations is based on the optimization of reproductive processes as a means of compensating for the loss of individuals through mortality. One way of reaching this equilibrium is the coadaptation of reproductive traits defining different strategies. I explored these tactics in lacertid lizards by analysing the covariation between, on the one hand, clutch size and frequency, and on the other, egg mass, using independent contrast data from 42 species. In addition, I examined the influence of female and newborn size on these variables. All the traits investigated, with the exception of clutch frequency, are influenced by female body size, reflecting physical constraints on reproductive output. The negative trade-off between clutch size and newborn size on one hand and egg mass on the other is congruent with the partitioning of energetic resources to produce a few large or numerous small descendents. Clutch frequency is unrelated to the other female reproductive traits.

The reproductive cycle of Lacerta agilis was studied in the Pyrenees at the southwestern boundary of the species range. The activity period is short (about 4.5 months). Males attain sexual maturity at smaller sizes than females and show mixed type spermatogenesis and spring spermiogenesis. Vitellogenesis in females is rapid (45 days for the whole population) with a long period of oviductal retention. Clutches are laid at the beginning of summer. During the reproductive period males mobilise reserves during the mating activities. Females deplete more fat reserves but later in the season than males and they are at their lowest after egg production. In both sexes, energy resources are mobilised from the fat bodies and also from the liver and the proximal part of the tail. Clutch size increases with female size but also depends on other factors. The short activity period does not appear to affect the reproductive cycle of this species. The gonad cycles and clutch traits do not show substantial differences from other European populations, perhaps with the exception of slight variation in the timing of reproductive events and, to a lesser extent, clutch frequency. This evidence strongly suggests that the limited southwestern range of Lacerta agilis is not due to environmental constraints but to very recent colonization.

Amat, F. & Llorente, G.A. & Carretero, M.A. (2003) -

A preliminary study of thermal ecology, activity times and microhabitat use of Lacerta agilis (Squamata: Lacertidae) in the Pyrenees. - Folia Zoologica, Prague, 52 (4): 413-422.

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Abstract. The thermal relations, diel, and annual activity and microhabitat use of Lacerta
agiliswere studied in the Pyrenean isolated range of this species. Although, this heliothermic
lizard demonstrated thermoregulatory capability, thermal constraints attributable to the mountain
climate were observed. The activity pattern was that typical of cold temperate lizards but the
beginning of activity was delayed in comparison with lowland populations. The reproductive
cycle determined different activity patterns for males and females. The ontogenetic and seasonal
changes in microhabitat use detected suggest influence of body size, reproductive condition in
adults, and interference with other individuals in juveniles.

We report the first study of diet composition of a Pyrenean lizard of the genus Iberolacerta, the Aurelio`s lizard, I. aurelioi, living in an alpine rocky slope at 2300 m of altitude in Andorra. Diet composition was studied during two years from 289 faecal pellets containing 966 prey items. Number of prey per pellet shows annual, seasonal and sexual differences. Diet of the species is poorly diverse, mainly dominated by insects, but we find clear evidences of opportunistic cannibalism based on the presence of toes and scales in feces. In addition, we detected minor sexual, annual and month differences in prey presence and abundance. Low amplitude of trophic niche width and prey diversity, as well as predation on clumped prey and cannibalistic behaviour suggest that Iberolacerta aurelioi has a remarkable dietary strategy in response to strong thermal and food availability constraints at high mountain habitats.

Lesotho Annotated Bibliography: Section 166: Reptiles including Annotated Species Checklist. Third Edition. - House 9 Publications, National University of Lesotho.

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Amer, S.A. & Reda, E.N. & Reheem, A.A.G.A. & El-Said, D.L. (2008) -

Lizard community and the genetic variability for two lacertid species in Damietta, North Egypt. - Egyptian Journal of Experimental Biology. (Zoology), 4: 229-235.

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The structure of the lizard community in Damietta was analyzed in the present study. Five locations were selected as a study area, each represents different habitat of Damietta Governorate. Seven lizard species were recorded which are related to four families and five genera. They were Chamaeleo africanus, Hemidactylus turcicus, Chalcides ocellatus, Mabuya quinquetaeniata, Mabuya vittata, Acanthodactylus boskianus and Acanthodactylus scutellatus. The lacertid lizards Acanthodactylus boskianus and Acanthodactylus scutellatus were the most common species in the community. Therefore, the study was extended to reveal the genetic variability for the population of these two species in Damietta. Four arbitrary chosen enzymes were analyzed electrophoretically. These enzymes were Glucose-6-phosphate dehydrogenase (G6PDH), esterase (EST), lactate dehydrogenase (LDH) and acid phosphatase (ACP). Eleven presumed gene loci have been recorded. A. scutellatus showed higher genetic variability with a polymorphism of 90% of its population compared to 82% for the samples of A. boskianus.

Prevalence and intensity of blood and intestinal parasites in a field population of a Mediterranean lizard, Lacerta lepida. - Parasitology Research, Berlin, 96 (6): 413-417.

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We describe the blood and intestinal parasites in the Ocellated lizard, Lacerta lepida, examining the factors that determine the prevalence and intensity of infection of haemogregarines, and the prevalence of coccidia and nematodes. In relation to haemogregarines, no juveniles were detected as being infected, whereas 71.7 % of adults were infected. The prevalence of infection was positively related to the size of the adults. There were no differences between seasons or sexes in the prevalence or intensity of infection in adults. There were no significant differences in the prevalence of infection by nematodes between ages or sexes, nor in relation to the size of adult lizards, but adult lizards excreting coccidian oocysts tend to be smaller. During the mating period, reproductive activities lead to a decrease in the body condition. However, neither the intensity of haemogregarine’s infection nor the prevalence of intestinal parasites was related to the lizards’ body condition.

Animals should tend to adjust the magnitude and characteristics of their escape responses according to the perceived levels of predation risk to cope with risk without incurring excessive costs. We analyze in the field the factors that determine the choice of escape behavior and patterns of refuge use of Wall Lizards under two simulated levels of predation risk and under variable environmental conditions, which may affect risk perception and costs of refuge use. The results show that Wall Lizards adjusted their antipredatory response according to several factors. The threat of predation posed by the predator affected the initial type of response of lizards but not the subsequent escape strategies employed. The escape strategy depended on the vulnerability to be captured (i.e., height on the wall and air temperature) and costs of refuge use (temperature and potential predation by ambush snakes). The initial risk of predation and thermal costs of refuge use affected emergence times from the refuge. The antipredator decisions of Wall Lizards, therefore, were influenced not only by the probability of mortality in the immediate future, such as the initial threat of predation and perceived susceptibility but also by consequences for long-term expected fitness, such as physiological costs of refuge use, and by the eventual risk of mortality associated with the use of unsafe refuges.

The threat sensitivity hypothesis assumes that multiple cues from a predator should contribute in an additive way to determine the degree of risk-sensitive behaviour. The ability to use multiple cues in assessing the current level of predation risk should be especially important to prey exposed to multiple predators. Wall lizards, Podarcis muralis, respond to predatory attacks from birds or mammals by hiding inside rock crevices, where they may encounter another predator, the smooth snake, Coronella austriaca. We investigated in the laboratory whether chemical cues may be important to wall lizards for detection of snakes. The greater tongue-flick rate and shorter latency to first tongue-flick in response to predator scents indicated that lizards were able to detect the snakes` chemical cues. We also investigated the use of different predatory cues by lizards when detecting the presence of snakes within refuges. We simulated successive predator attacks and compared the propensity of lizards to enter the refuge and time spent within it for predator-free refuges, refuges containing either only visual or chemical cues of a snake, or a combination of these. The antipredatory response of lizards was greater when they were exposed to both visual and chemical cues than when only one cue was presented, supporting the threat sensitivity hypothesis. This ability may improve the accuracy of assessments of the current level of predation risk inside the refuge. It could be especially important in allowing lizards to cope with threats posed by two types of predators requiring conflicting prey defences.

Prey often respond to predators by increasing refuge use, but some refuges may expose prey to other types of predators. Because predators are not always visible inside refuges, their chemical stimuli may provide early warning of their presence. However, in ectotherms, chemosensory detection of predators may be thermally dependent. Lizards often hide in cold refuges, where their body temperature (Tb) may decrease, and this might affect their ability to detect chemicals of saurophagous snakes that ambush inside refuges. We examined the effects of Tb of wall lizards, Podarcis muralis, on their ability to detect chemicals of smooth snakes, Coronella austriaca. Differential tongue flick (TF) rates and behavioral patterns of lizards in response to scent of smooth snakes showed that lizards with optimal Tb discriminated and avoided the snake`s scent, whereas lizards with suboptimal Tb did not. We also examined the importance of this effect on movement patterns and refuge use by lizards in terraria with safe (odorless) or unsafe refuges (snake-scented). While results of this experiment were variable, there was evidence of snake avoidance in that lizards at optimal Tb spent less time in snake-scented refuges relative to odorless refuges. Therefore, this study provides evidence that chemosensory discrimination of snake chemicals is thermally dependent, and, thus, suboptimal Tb impedes a lizard`s ability to avoid snake-scented refuges.

Many animals use chemical cues to detect predators. However, according to the threat sensitivity hypothesis, natural selection should favor individuals able to respond differentially to the chemical cues of predators that pose different levels of risk rather than responding to chemical cues of all predators in a similar way. The behavioral responses of prey to chemical cues of predators might depend, for example, on the degree to which predators include prey in their diets. The Wall Lizard (Podarcis muralis) is under high predation pressure by saurophagous-specialist Smooth Snakes (Coronella austriaca). Wall Lizards are also an important part of the diet of the Montpellier Snake (Malpolon monspessulanus) and occasional prey of the Ladder Snake (Elaphe scalaris). Also, Wall Lizards are sympatric with the non-saurophagous Viperine Snake (Natrix maura). In the laboratory, we compared the tongue-flick rates and behavioral patterns of Wall Lizards in response to exposure to chemical signals of these four snake species that pose different risks of predation. Results suggested that Wall Lizards were able to discriminate the chemical cues of the predatory snakes from the non-saurophagous snake. However, lizards did not respond differentially to chemical cues of different predatory snakes. We suggest that other additional cues are probably needed to elicit an adequate, specific defensive response.

Amo, L. & López, P. & Martin, J. (2004) -

Prevalence and intensity of Haemogregarinid blood parasites in a population of the Iberian Rock Lizard, Lacerta monticola. - Parasitology Research, Berlin, 94: 290-293.

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The study of the effect of parasites on their host populations is essential for understanding their role in host population dynamics and ecology. We describe the general field population biology of haemogregarines in the Iberian rock lizard, Lacerta monticola, examining the factors that determine the prevalence and intensity of infection. Prevalence and infection intensity were higher in adults than in juvenile lizards. The prevalence rate was higher in larger lizards, probably because they were also older and had been more often in contact with parasites during their lifetime. During the mating season, the intensity of parasite infection was greater in males than in females, probably due to immunosuppressive effects of testosterone. The parasite load had a negative effect on the body condition during the reproductive season. The results suggest that the interactions between parasites and hosts are not stable in this lizard population.

Multiple predators can have risk-enhancing effects due to conflicting predator-specific prey defenses. Wall lizards (Podarcis muralis) escape from searching predators by fleeing into refuges where they can also encounter ambush snakes (Coronella austri- aca). We simulated attacks to lizards in the laboratory to compare their use of preda- tor-scented refuges with that of control refuges, and examined whether lizards that suf- fered a second attack changed the refuge used in the first attack if they had been hidden before in an unsafe refuge. The presence of snake cues neither influenced time to enter the refuge nor refuge choice. However, appearance time was greater in the snake- scented refuge, probably due to time spent acquiring information to ensure the absence of the snake. This was supported because once lizards appeared, they resumed their activities in the same interval of time. Our results suggest a case of predator facilitation due to conflicting prey defenses.

Animals in natural environments usually face different types of predators, and conflict- ing prey defenses can increase risk of predation if prey responses to one predator result in a greater risk from another predator. Wall lizards (Podarcis muralis) respond to predators in the open, such as birds or mammals, by hiding inside rock crevices, where they may be exposed to predation by ambushing saurophagous smooth snakes (Coro- nella austriaca). This offers an excellent system to study how prey behave to avoid multiple predators acting simultaneously. We first performed a field study to examine the escape behavior and refuge use of wall lizards. Lizards mainly responded to the simulated attack by fleeing to hide in the nearest refuge. However, lizards that were far from known refuges sometimes used an alternative escape behavior; lizards fled for longer while passing potential refuges without hiding, thus, presumably avoiding hiding when their reliance in refuge safety was low. This was supported by a further analysis of movement patterns of wall lizards. Spontaneous locomotor patterns could be described as an amount of bursts of locomotion separated by short and long pauses. Lizards stopped near refuges more often, but previously they spent more time in short pauses when moving in or close to refuges exploring them by tongue flicking, than in open rocks. This agreed with a laboratory experiment where we analyzed responses of lizards to substrate scent deposits of smooth snakes. Lizards detected the snake’s scent, and responded by increasing their movement rate in an effort to leave the risky area. However, in the field, after a simulated attack in the open, lizards emerged from the refuge and left the area quickly, decreasing the frequency of short pauses, probably to avoid a new attack in the open. Flexibility in antipredatory behavior may help wall lizards to avoid the risk enhancing effect of two types of predators requiring conflict- ing prey defenses.

We describe the general field population biology of haemogregarines and mites in the wall lizard, Podarcis muralis, examining the factors that determine the prevalence and intensity of infection. The intensity of infection by haemogregarines in females was slightly lower in summer than in spring, whereas males maintained similar levels of intensity among all the seasons, probably due to immunosuppressive effects of testosterone. This is also supported because the prevalence and the infection intensity by mites were higher in males than in females. Parasite load was higher in lizards with better body condition, which could be reflecting the mortality of infected lizards with poor condition.

Amo, L. & López, P. & Martín, J. (2005) -

Chemical Assessment of Predation Risk in the Wall Lizard, Podarcis muralis, is Influenced by Time Exposed to Chemical Cues of Ambush Snakes. - The Herpetological Journal, 15 (1): 21-25.

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Lizards often respond to predator presence by increasing refuge use. However, this behaviour may expose lizards to saurophagous snakes, which inhabit the same refuges to ambush their lizard prey. Snakes, which are not always visible, deposit chemical trails that can be detected by lizards. Even though there are obvious advantages of using chemical cues, chemical detection of predators might lead to very conservative estimates of risk. This is because chemical cues might indicate that an area was risky in the recent past, but not necessarily at the current time. We examined experimentally whether wall lizards (Podarcis muralis) avoid using refuges that contain chemical cues of smooth snakes (Coronella austriaca), and whether this avoidance response is maintained long term or whether it can be modified. Results suggest that wall lizards detected the chemical cues of smooth snakes inside refuges, and, in the short term, decreased the use of predator-scented refuges and increased their escape movements. However, this avoidance response seemed to decrease in the long term. By investigating the refuge again over subsequent time periods, lizards reassessed whether the snake was actually present, modified their refuge use and decreased their avoidance response. Therefore, wall lizards seem able to assess temporal variations in predation risk by snakes inside refuges and to respond accordingly.

Amo, L. & Lopez, P. & Martin, J. (2006) -

Nature-based tourism as a form of predation risk affects body condition and health state of Podarcis muralis lizards. - Biological Conservation, 131: 402-409.

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Nature-based tourism has experienced a greater increase, and, even although it might have deleterious effects for many wildlife species, its consequences remains little explored. Many lizard species are endangered and tourism has been proposed as a potential factor responsible of the decline of several lizards` populations, but no study has examined the effect of tourism on lizards` behaviour, body condition and health state. Many lizards respond to people as if they were predators, by readily escaping to refuges. However, an increase in the frequency of these antipredatory strategies can lead to a loss of body condition, which may have important consequences for short and long term fitness. We analysed the effects of tourism on escape behaviour of common wall lizards, Podarcis muralis, as well as on their body condition and health state (ectoparasites, blood parasites, and cell mediated immune response). Results showed that lizards did not modify their escape behaviour in response to tourism. Lizards had similar approach and flight distances, and escape strategies regardless of the level of tourism pressure. However, lizards inhabiting areas with high tourism levels, where they presumably needed to perform antipredatory behaviours more often, showed higher intensity of infection by ticks and lower body condition at the end of the breeding period. Moreover, lizards with poorer body condition had lower cell mediated immune responses. Therefore, tourism seems to have deleterious effects on body condition and on host-parasite relationships in this lizard species. These effects should be taking into account when designing walking paths in protected areas. Our study reports evidence that regardless lizards showed similar escape behaviour in tourist than in natural areas, their body condition and health state should be also examined to accurately assess the actual effects of tourism on lizards` populations.

The ability to use multiple cues in assessing predation risk is especially important to prey animals exposed to multiple predators. Wall lizards, Podarcis muralis, respond to predatory attacks from birds in the open by hiding inside rock crevices, where they may encounter saurophagous ambush smooth snakes. Lizards should avoid refuges with these snakes, but in refuges lizards can also find non-saurophagous viperine snakes, which lizards do not need to avoid. We investigated in the laboratory whether wall lizards used different predator cues to detect and discriminate between snake species within refuges. We simulated predatory attacks in the open to lizards, and compared their refuge use, and the variation in the responses after a repeated attack, between predator-free refuges and refuges containing visual, chemical, or visual and chemical cues of saurophagous or non-saurophagous snakes. Time to enter a refuge was not influenced by potential risk inside the refuge. In contrast, in a successive second attack, lizards sought cover faster and tended to increase time spent hidden in the refuge. This suggests a case of predator facilitation because persistent predators in the open may force lizards to hide faster and for longer in hazardous refuges. However, after hiding, lizards spent less time in refuges with both chemical and visual cues of snakes, or with chemical cues alone, than in predator-free refuges or in refuges with snake visual cues alone, but there were no differences in response to the two snake species. Therefore, lizards could be overestimating predation risk inside refuges. We discuss which selection pressures might explain this lack of discrimination of predatory from similar non-predatory snakes.

Natural vegetation in Europe appears nowadays deeply modified by human activities. In the Guadarrama Mountains (Central Spain), ancient reforestations with Scots pines, Pinus sylvestris, replaced original deciduous pyrenean oak, Quercus pyrenaica, forests (since the Roman period). However, the effect of reforestations on fauna remains little known, especially in reptiles. We described patterns of microhabitat selection in several species of Lacertid lizards, and analyzed whether the modification of the original vegetation affected distribution and population densities of lizards. The species of lacertid lizards found in oak forests (Psammodromus algirus, Lacerta lepida and Podarcis hispanica) were different to those of in pine plantations (Podarcis muralis and Podarcis hispanica). Lizards did not use habitat at random and this could explain differences in species found in both forests, which differed in some microhabitat structure characteristics. Most lizards selected microhabitats with rocky outcrops, with low cover of trees, and close to refuges. These microhabitat preferences also explained abundance of lizards in transects. From the perspective of conservation and management of lizards, pine plantations seem not to contribute too much to the diversity of lizard species because species typical from oak forests were lost. This study has implications for pine reforestation management, because allowing the recolonization by understory oaks, and leaving some open areas, without trees but with dense shrubs and rocks inside reforestations would contributed to maintain lizard populations.

We analyzed whether habitat deterioration due to ski slopes affected lizards’ behavior, and whether these changes in behavior had consequences for the body condition and health state of lizards. Results suggested that habitat deterioration in ski slopes not only implied a loss of optimal habitat for lizards, but also led to an increase in perceived risk of predation. Males seemed to adjust their movement patterns to differences in risk, increasing their movement speed during their displacements across risky areas within ski slopes, but as a consequence, they incurred loss of body condition. A laboratory experiment supported that fleeing at high speeds in areas without refuges can be a factor responsible for reduced body condition. However, changes in body condition did not affect sprint speed of lizards. Our study provides new evidence that behavioral strategies to cope with increased predation risk, due to human-induced habitat deterioration, may affect body condition of lizards. Our results have applications for the design of conservation plans for this endangered lizard species. Environmental impact studies should be performed before opening new ski slopes, but taking into account not only the presence of lizards but also their body condition. An effective way to decrease the effects of habitat loss may be the artificial restoration of some refuges to create safe corridors for movement.

Lizards often respond to increased predation risk by increasing refuge use, but this strategy may entail a loss of thermoregulatory opportunities, which may lead to a loss of body condition. This may be especially important for pregnant oviparous female lizards, because they need to maintain optimal body temperatures as long as possible to maximize developmental embryos rate until laying. However, little is known about how increased time spent at low temperatures in refuges affects body condition and health state of pregnant female lizards. Furthermore, it is not clear how initial body condition affects refuge use. Female Iberian rock lizards forced to increase time spent at low temperatures showed lower body condition and tended to show lower cell-mediated immune responses than control females. Therefore, the loss of thermoregulatory opportunities seems to be an important cost for pregnant females. Nevertheless, thereafter, when we simulated two repeated predatory attacks, females modified refuge use in relation to their body condition, with females with worse condition decreasing time hidden after attacks. In conclusion, female lizards seemed able to compensate increased predation risk with flexible antipredatory strategies, thus minimizing costs for body condition and health state.

Amr, Z.S. & Al-Oran, R. & Disi, A.M. (1994) -

Reptiles of southern Jordan. - The Snake, 26: 41-49.

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A total of 46 reptilian species belonging to 9 families (Gekkonidae, Agamidae, Chamaeleonidae, Scincidae, Lacertidae, Varanidae, Boidae, Colubridae and Viperidae) are reported from Southern Jordan. Bunopus tuberculatus, Tropiocolotes steudneri and Phrynocephalus arabicus are new records, while Coluber elegantissimus is reported for the second time. A. population of horned Cerastes cerastes was found at Disah area. Remarks on the distribution of these reptiles are included.
Acanthodactylus boskianus, Acanthodactylus tristrami, Acanthodactylus grandis, Acanthodactylus opheodurus, Acanthodactylus pardalis, Ophisops elegans, Mesalina guttulata, Lacerta laevis kulzeri.

We present the results of several field campaigns, realized from June 2007 to September 2008, and from March 2012 to May 2014 in different Algerian localities (Cap Djinet, Tigzirt, Boukhalfa, Ouadias, El Misser, Darna and Talaguilef). During this field work we have collected 488 specimens belonging to 35 species: 328 small mammals (11 species), 53 amphibians (6 species), 106 squamates (17 species) and one chelonian. The trapping success was highly variable (0.1 to 13.34 %). Genetic and morphometric analyses in progress will improve our knowledge of the evolutionary history of these small vertebrates at population and community scales.

This paper reports the results of newest investigations of lizards` demonstration behaviour. A special attention was paid to lizards` aposematic and epigamious reactions which are of great importance when forming couples and defending individual territories.
The species differences represent in quite a number of cases the most essential obsacle for the formation of couples of the individuals of different specis, even if the morphological differences of these species are hardly perceptible. Being the determining feature in intra- and inter- species contacts, the demonstration behaviour ensures the integrity of a species.

Morphometric analysis of limb proportions of five sympatric species of desert lizards (Sauria, Eremias) in the southern Balkhash Lake region. - Proceedings of the Zoological Institute of the Academy of Science (U.S.S.R.), 74: 3-13.

Morphometric evaluation of limb proportions in five sympatric desert lacertas of the genus Eremias, namely E. velox, E. grammica, E. intermedia, E. lineolata, and E. scripta inhabiting southern Balkhash Lake region (Sary-Ishyk-Otrau desert), Kazakhstan, was made. Considerable species differences in proportions of some limb segments associated with locomotion adaptations (burrowing, climbing etc) of these lizards have been found as well as sexual dimorphism and age variability. Associations among morphometric peculiarities (limb proportions), locomotion activity, spatial, temporal, and trophic characteristics of ecological niches of sympatric species are discussed.

Morphometric evaluation of limb proportions in five sympatric desert lacertas of the genus Eremias, namely E. velox, E. grammica, E. intermedia, E. lineolata, and E. scripta inhabiting southern Balkhash Lake region (Sary-Ishyk-Otrau desert), Kazakhstan, was made. Considerable species differences in proportions of some limb segments associated with locomotion adaptations (burrowing, climbing etc) of these lizards have been found as well as sexual dimorphism and age variability. Associations among morphometric peculiarities (limb proportions), locomotion activity, spatial, temporal, and trophic characteristics of ecological niches of sympatric species are discussed.

TERRA TYPICA. Settlement Terskol on the Baksan River, Kabardino-
Balkaria, Russia.
DISTRIBUTION. Endemic of the high-mountainous Caucasus. It
occurs in the high-altitude part of the western half of the Great
Caucasus Range – from Elbrus in the east to the mountains Fisht
and Osten in the west. The eastern border of the distribution
range runs in the upper reaches of Baksan River on the south-eastern
slopes of Elbrus and mountain Cheget in Kabardino-Balkaria.
Further to the west it is widely distributed in the upper reaches of
the rivers Kuban’, Teberda, Zelenchuck, Laba, Belaya and others
flowing down to the north. It inhabits the southern slopes of the
Caucasus: in the upper reaches of the Bzyb River and its tributaries
in Abkhazia and the upper reaches of Mzymta River in the southeast
of the Krasnodar Territory.
CONSERVATION STATUS. The number is rather stable everywhere
within of its distribution range. The species is included into the
Red Data Book of IUCN with the status DD.

TERRA TYPICA. Settlement Yelenovka (= Sevan) on the shore of
the Sevan Lake, Armenia.
DISTRIBUTION. It is widely distributed in the mountains of northwestern
and northern Armenia and adjacent regions of southern
Georgia and western Azerbaijan. Outside of the Caucasus it is
also known from the north-western Turkey. In 1967 D. armeniaca
was introduced into the Ukraine. At present this parthenogenetic
species formes a stable extensive population in the rock canyon
of Teterev River of the Zhitomir region.
CONSERVATION STATUS. The number is high and stable everywhere
within the whole distribution range.

TERRA TYPICA. Settlement Krasnaya Polyana in the south of the
Krasnodar Territory, Russia.
DISTRIBUTION. Endemic of the montane Caucasus. It is widely
distributed in the western part of the Great Caucasus Range, on
northern, western and south-western slopes of the Black Sea
chain of the Great Range in the Krasnodar Territory, Abkhazia and
north-western and south-western Georgia, to the canyons of the
rivers Kodory and Ingury in lower and upper Svanetia in the east.
The northern border of its distribution passes in the area of the
middle current of numerous left tributaries of the Kuban River. The
most northern habitats of the species are known in the vicinity of
Goryachyi Klyuch on Psekupsa River and the vicinity of the town
Maikop. The southern border of the distribution range stretches
on the southern slopes of the Great Range and, gradually lowing
down, on the western spurs of the Gagra range. It comes out
to the sea in the south of the Krasnodar Territory and western
Abkhazia, where it is sporadically recorded on the coast from the
ALPINE LIZARD DAREVSKIA ALPINA (DAREVSKY, 1967)
ARMENIAN LIZARD DAREVSKIA ARMENIACA (MEHELY, 1909)
BRAUNER’S ROCK LIZARD DAREVSKIA BRAUNERI (MEHELY, 1909)
81
L A C E R T I D A E
town Anapa in the north up to the canyons of the rivers Khoby
and Riony in the south-west of Georgia.
The nominative subspecies D. b. brauneri occurs in the south
of the Krasnodar Territory, in Abkhazia and adjacent regions of
Georgia. D. b. darevskii (Szczerbak, 1962) inhabits foothills of
the western part of the Great Caucasus range, mainly within the
Krasnodar Territory. A comparatively small range of the subspecies
D. b. szczerbaki (Lukina, 1963) occupies a narrow strip of
coastal rocky steeps, sporadically occurring along the coast of the
Black Sea from the town Anapa in the north and for more than
100 km further to the south-east towards Novorossiisk.
CONSERVATION STATUS. On the most part of the distribution range
the number of the species is rather high and stable.

TERRA TYPICA. Kazbek Mountain, Georgia.
DISTRIBUTION. Endemic of the Great Caucasus. It is widely distributed
on the northern and southern slopes of the Great Caucasus
Range from the northern foothills of Elbrus in Kabardino-Balkaria
in the west up to the south-eastern extreme point of its Caspian
chain within Azerbaijan in the east. South-western border of the
distribution range passes on the southern foothills of Megrel’sky,
Svanetsky and Lekhchumsky mountain ranges in lower Svanetia,
and then coming down within South Ossetia approximately up to
the latitude of the health-resort Dzhava. Along the southern spurs
of Kharul’sky, Kartalinsky and Kakhetinsky mountain ranges it comes
out into the slopes of the Water Dividing (Watershed) Range in the
Alazan River valley of Georgia. In Azerbaijan this border continues
in the upper reaches of the rivers beginning from the Great Range,
and further in the east – from the slopes of the mountains Baba-
Dag and Dyubrar, in particular of the rivers Pirsagat and Sumgait.
This lizard is also widely distributed on the northern slopes of the
Great Caucasus, from the Baksan ravine in Kabardino-Balkaria in
the west to Samur range in the South Dagestan in the east.
The north-western border of the distribution area passes
here in the Baksan ravine approximately from the town Tegenekly
in the south, continuing then to the east on the northern foothills
of the Skalisty and Bokovoi ranges. There the most northern
habitats are known from the ravine Adyr-su, near the waterfall
in the Chegemskaya gorge, from the vicinity of the Blue Lakes in
Kabardino-Balkaria and the vicinity of Alagir and the settlement
Balta in the lower part of the Dar’yal ravine of North Ossetia. In
the foothills and on the northern slopes of the Andiysky mountain
range in the south-eastern Chechnya and possibly in the adjacent
part of Ingooshetia occurs a narrow-ranged subspecies D. c. vedenica
(Darevsky et Roitberg, 1999).
CONSERVATION STATUS. The number of the species everywhere
within its distribution range is high and stable.

TERRA TYPICA. Town Enzely, Iran, southern coast of the Caspian
Sea.
DISTRIBUTION. It inhabits mainly forested northern and northwestern
foothills of the Elburz range in northern Iran and Lenkoran
lowland in the south-eastern foothills of the Talysh range in the
south-east of Azerbaijan. There the most northern habitats are
known from the valley of Vilyazhchai River.
CONSERVATION STATUS. On the most part of the distribution range
the number of the species is high and stable.

TERRA TYPICA. 20 km to the west from the settlement Borchka,
vilajet Artvin, north-eastern Turkey.
DISTRIBUTION. The main distribution range of the species occupies
the northern spurs of the East-Pontic mountains in the
north-east of Turkey, at least up to the vilajet Giresun in the west.
A small isolated area of the distribution range is known on the
western slopes of the mountain Mtirala in the vicinity of the town
Batumi in the south-west of Ajaria (Georgia).
CONSERVATION STATUS. As a rare species the species was included
into the Red Data Book of the USSR (1984) – category 3 and the
Red Data Book of IUCN with the status ENC2a.

TERRA TYPICA. Settlement Levashi in the region with the same
name of central Dagestan.
DISTRIBUTION. It occurs in the montane Dagestan, southern
Chechnya, Ingooshetia and south-east of north Ossetia, from the
lower part of the Dar’yal ravine in the west up to Gimrinsky range
in the east. The northern border of the distribution area passes in
the foothills of the Skalisty range in the upper current of numerous
right tributaries of Sunja River. Further on the extreme north-eastern
foothills of the frontal ranges of Dagestan it attains the nearest
vicinity of Makhachkala city in the east. An isolated population
lives also on the southern slope of the Great Caucasus range in
the canyon of the river Bol’shaya Liakhvy on the territory of South
Ossetia in Georgia.
CONSERVATION STATUS. On the most part of the distribution range
the number is stable.

TERRA TYPICA. Settlement Shagaly, Armenia.
DISTRIBUTION. It is comparatively widely distributed in the north
Armenia and southern Georgia up to the foothills area in the valley
of Kura River in the north. The recent distribution range is divided
into some isolated populations.
CONSERVATION STATUS. On the most part of the distribution range
the number is stable.

TERRA TYPICA. Valley of Lar, to the north-west of Teheran, northern
Iran.
DISTRIBUTION. The distribution range of the species covers practically
all Alborz range in the north of Iran and adjacent east Kopet
Dagh range both in its Iranian part and in the south of the neighbouring
Turkmenistan. In Turkmenistan the species is very rare and
once it was considered to be extinct. In central Kopet Dagh it is
known here from a few points in the ravines Sushanka and Bol’shie
Karanki.
CONSERVATION STATUS. In Iran the number of the species is high
and stable. In Turkmenistan the number is catastrophically declining.
The species is included into the Red Data Book of Turkmenistan
(1999): category 3 – a rare species on the periphery of the distribution
range.

TERRA TYPICA. Vicinity of the town Artvin, north-eastern Turkey.
DISTRIBUTION. South-west of the Krasnodar Territory, western
and north-eastern Georgia and the extreme north-west of
Azerbaijan. Outside of the Caucasus it occurs on the extreme
north-east of Turkey. Of 6 described subspecies 5 are distributed
in the Caucasus. D. d. silvatica (Bartenjev et Reznikova, 1931) inhabits
submontane part of the Krasnodar Territory mainly in the
Caucasian Biosphere State Reserve and the adjacent spurs of
the Great Caucasus. D. d. boehmei (Bischoff, 1982) lives in the
western spurs of the Great Caucasus in Krasnodar Territory and
adjoining regions of Abkhazia up to the valley of the river Kodor in
the south-east. D. d. abchasica (Bischoff, 1982) inhabits the coast
of the Black Sea of Abkhazia and the adjacent regions of western
Georgia, D. d. barani (Bischoff, 1982) is known from coastal mountain
regions of Ajaria and of neighbouring part of north-eastern
Turkey. D. d. orlowae (Bischoff, 1984) inhabits southern slopes
and foothills of the Great Caucasus in Georgia and north-western
Azerbaijan.
CONSERVATION STATUS. The number is rather high and stable.

TERRA TYPICA. Ravine of small Charnali River in the lower reaches
of the river Chorokh in the extreme south-west of Ajaria (southwestern
Georgia) and the adjacent part of Turkey.
DISTRIBUTION. It occurs on a limited territory in the north-eastern
foothills of the Pontic range, within extreme south-west of Ajaria in
Georgia and adjoining regions of north-eastern Turkey. The distribution
is not sufficiently studied.
CONSERVATION STATUS. The number is rather high and stable.

TERRA TYPICA. The Crimean Peninsula.
DISTRIBUTION. The distribution range of the species is limited by
the mountain Crimea from Sevastopol in the west up to the cape
Kiik-Atlam near the town Feodosia in the east. The northern border
of the distribution range runs on the northern slopes of the
Crimean Mountains through the towns Bakhchisarai, Simferopol
and the vicinity of the settlement Vishennoe in the Belgorod region.
A small isolated population lives on the rocks Adalary near
Gurzuf.
CONSERVATION STATUS. The number is rather high and stable.

TERRA TYPICA. Town Abastumani in south-western Georgia.
DISTRIBUTION. Endemic of Georgia. The recent distribution range
forms some more or less extensive isolates mainly situated on the
slopes of the Meskhetsky range within Ajaria and western Georgia
up to Borzhomi ravine in the valley of Kura River and north-western
slopes of Trialetsky range in the east. Isolated populations live
on the southern slopes of Lechkhumsky and Rachinsky ranges in
the foothills of Great Caucasus, where this lizard is known from the
vicinity of the towns Kutaisi, Tkvibuli and the health-resort Labarde.
Its distribution in the foothills of the Great Caucasus Range is still
not sufficiently studied.
CONSERVATION STATUS. The number in the most of known populations
is rather high and stable.

TERRA TYPICA. Town Artvin, north-eastern Turkey.
DISTRIBUTION. The distribution range of the species covers
north-eastern Turkey (approximately from Erzerum in the west and
from the valley of the upper current of Aras River in the south) and
northern and southern foothills and spurs of the Meskhetsky range
within Ajaria and neighbouring regions of Georgia from the coast
of the Black Sea in the west up to the middle current of Kura River
in the east. There are local isolated populations in the ravines of
several lateral tributaries of the river Kura (Akhaldaba, Baniskhevi,
Azkuri and others). In Georgia and in the ravine of the lower current
of river Chorokh in Turkey occurs subspecies D. p. adjarica
(Darevsky et Eiselt, 1980). The remaining part of the distribution
range is occupied by the nominative subspecies D. p. parvula.
CONSERVATION STATUS. On the most part of the distribution range
the number of the species is high and stable.

TERRA TYPICA. City Tiflis (= Tbilisi), Georgia.
DISTRIBUTION. The main distribution range of the species occupies
right-bank section of the valley of middle current of Kura
River and the ravine of its right tributaries within Georgia, northern
Armenia and north-western Azerbaijan. It covers the area from
Gori in the west up to northern foothills of the ranges of the Lesser
Caucasus in the lower and middle current of the rivers flowing
into Kura River: Algeti, Khrami, Debeta, Akstafa-chai, Gyandzhachai
and others in the south-east. Big population, isolated now
from the main distribution range, is known in the valley of middle
current of the left tributary of Kura River – river Iori on the southern
slopes of the Zivgamborsky range. A great zoogeographical
interest represents an isolated population of this species discovered
in south-western Azerbaijan, in the ravine of upper current
of Akar River, belonging to the basin of Aras River. Presence of D.
portschinskii in the southern foothills of Lesser Caucasus can be
connected with a radical reconstruction of river systems which
was taking place on the territory of Transcaucasia in the postglacial
period. The nominative subspecies D. p. portschinskii occupies
the most part of the distribution range. In the ravines of right tributaries
of Kura River – the rivers Mashavera in Georgia and Debeta
in Armenia – occurs D. p. nigrita (Bakradze, 1976).
CONSERVATION STATUS. On the most part of the distribution range
the number is high and stable.

TERRA TYPICA. Settlement Narzan, North Caucasus.
DISTRIBUTION. It occurs in the north-west of the Balkan Peninsula
from one side and within the Caucasus and adjacent part of Iran
from the other side. It is sporadically distributed on the both sides
of the Great Caucasus Range within the limits of the Krasnodar
and Stavropol Territories, in Kabardino-Balkaria, North Ossetia,
Chechnya, Ingooshetia and Dagestan, and in the Transcaucasia
– on the coast of the Black Sea of Abkhazia, in north Georgia,
northern Azerbaijan. Isolated part of range lies in south-east
Azerbaijan and in the north of Armenia. D. p. praticola inhabits
the most part of the distribution range. D. d. pontica (Lantz et
Cyren, 1919) inhabits the coast of the Black Sea and north-western
Caucasus.
CONSERVATION STATUS. On the most part of the distribution range
the number is high and stable.

TERRA TYPICA. Vicinity of the settlement Ijevan, Armenia.
DISTRIBUTION. It has a comparatively limited distribution range in
the northern Armenia and adjacent regions of Azerbaijan. The recent
distribution range is divided into several more or less isolated
populations. One of them, now extinct, existed on the north-eastern
shore of the Lake Sevan in Armenia.
CONSERVATION STATUS. The number is stable.

TERRA TYPICA. Town Trapezund (Trabzon), north-eastern
Turkey.
DISTRIBUTION. The species is widely distributed, mainly in
the western Transcaucasia, and in the north – in the montane
Ingooshetia, Chechnya, neighbouring Dagestan and in northern
Ossetiya. Outside of the Caucasus it is widely distributed in the
northern half of Turkey. Of eight described subspecies the five
occur in the Caucasus. D. r. bischoffi (Boehme et Budak, 1977)
occurs in Ajaria. D. r. obscura (Lantz et Cyren, 1936) inhabits the
valley and adjoining slopes of mountains of the upper current of
Kura river within Georgia. D. r. macromaculata (Darevsky, 1967)
is recorded in the ravine of the right tributary of Kura River, river
Akhalkalakichai in southern Georgia. D. r. svanetica (Darevsky et
Eiselt, 1980) occurs in the ravine of the upper current of the river
Inguri in Svanetia (western Georgia). D. r. chechenica (Eiselt et
Darevsky, 1991) is widely distributed on the southern slopes of
the Great Caucasus from lower Svanetia in the west up to northwestern
Azerbaijan in the east, and also isolatedly on the northern
slopes within the limits of the montane Chechnya and Dagestan.
CONSERVATION STATUS. The number of the species is high and
stable everywhere.

TERRA TYPICA. Town Kislovodsk, Stavropol Territory.
DISTRIBUTION. The main distribution range of the species
covers the northern slopes of the Rocky range within Karachaevo-
Cherkesskaya Autonomous Region, south of the Stavropol Territory
and Kabardino-Balkaria, from the valley of the upper current of
Kuban’ River and its tributaries in the west up to the ravine of the
middle current of the river Chegem in the east. The most northern
isolated population (which was considered as extinct) lives in the southern foothills of the Stavropol Upland, in the limits of
the Alexandrovsky region of the Stavropol Territory. The southern
border of the distribution range passes on the north-eastern
spurs of the mountain Elbrus in the ravine of the river Kuban, and,
crossing the Rocky Range (known from the Mariinsky pass), continues
further on its watershed in the upper reaches of the rivers
Kuma, Podkumok, Alikanovka, Beryozovaya, Zolka and others. On
the ravine of the river Baksan it comes up to the town Tyrny-Auza
and goes from the Chegem waterfalls up along the Chegem River
gorge.
CONSERVATION STATUS. The number is high and stable.

TERRA TYPICA. Settlement Akhta, ravine of the river Razdan,
northern Armenia.
DISTRIBUTION. The distribution range of the species is divided
into several isolated populations within the northern and central
high-mountain Armenia and the adjacent regions of eastern
Turkey, from where along the valley of Kura River this lizard penetrates
into southern Georgia. Outside of the Caucasus it occurs in
the valley of the river Aras in eastern Turkey.
CONSERVATION STATUS. The number is high and stable.

TERRA TYPICA. Town Bazarchai (Basarkent), Sisiansky region,
eastern Armenia.
DISTRIBUTION. The recent distribution range of the species is divided
into several of more or less extensive isolates, attributed to
the steppe high-mountain regions of the mountain-steppe zone
of Armenia, Azerbaijan, southern Georgia and eastern Turkey. The
most extensive areas of the distribution range are occupied by
the high-mountain zone of the Gegamsky range up to the shores
of Sevan Lake in the north, a ring-like high-mountain area of the
Aragaz mount and a high-mountain plateau in the foothills of the
Childyrsky and Javakhetsky ranges in the extreme north-west of
Armenia and adjoining regions of southern Georgia. Besides, it is
encountered on the Ardenissky range confining the basin of Sevan
Lake from the south. Further to the east it inhabits all high-mountain
part of the Karabakh Upland within the limits of Nagorny Karabakh.
Outside of the Caucasus, covering the mountains, surrounding the
Lake Van, the species is widely distributed in north-eastern, eastern
and south-eastern Turkey, at least from the range Palandeken
in the vilajet Erzerum in the west. Probably it penetrates into the
high-mountain region of extreme west on the territory of the
neighbouring Iran.
Three subspecies are described. The nominative subspecies
D. v. valentini is distributed in the whole Transcaucasia. D. v. lantzicyreni
(Darevsky et Eiselt, 1967) occurs on the most part of Turkey.
The third subspecies D. v. spitzenbergerae (Eiselt, Darevsky et
Schmidtler, 1992) is known only from the type locality in the vilajet
Khakyari in the extreme south-east of Turkey.
CONSERVATION STATUS. On the most part of the distribution area
the number of the species is high and stable.

TERRA TYPICA. China, Chifu (now Yantai, Shandun Province).
DISTRIBUTION. The species is widely distributed in the eastern
part of Mongolia, in China (from the lake Kukunor up to Shanghai
city), in the western part of the Korean Peninsula and in Russia,
in the southern Buryatia (to the north approximately up to Ulan-
Ude city) and in the extreme south-west of the Chita region. In
Mongolia the most part of the findings are attributed to the north
and east from the range Khangai in Kobdos, Gobi-Altai, Dzabkhan,
Bayan-Khongor, South-Gobi, Middle-Gobi, Central, Bulan, Selengin,
Khentei, East-Gobi, Sukhe-Bator and Eastern aimaks. Of two
known subspecies within North Eurasia occurs the western one
– E. a. barbouri Schmidt, 1925.
CONSERVATION STATUS. The species (E. a. barbouri) is included
into the Red Data Book of the Russian Federation (2001): category
4 – species uncertain in its status distributed in Russia on the
extreme border of its distribution range.

TERRA TYPICA. Interstream area of the rivers Ural and Emba.
DISTRIBUTION. The distribution range stretches from northeastern
Romania in the west to south-western Mongolia (Jungar
Gobi) and north-western China (Xinjiang-Uyghur Autonomous
Region and Inner Mongolia) in the east, Turkey and Iran in the
south. Within the limits of North Eurasia it is widely distributed
from the steppe zone of Moldova, the Ukraine and the European
part of Russia including North Caucasus and the area of the Volga river in the west, to Kazakhstan, the Republics of Middle Asia
and Mongolia in the east. In Mongolia records are known in the
southern part of the Kobdos aimak and in the Gobi-Altai aimak.
Disjuncted from the main distribution range the species occurs
in the eastern Transcaucasia, within the limits of Azerbaijan,
eastern Georgia and the basin of the Sevan Lake in Armenia.
The most northern, evidently isolated habitats are known in the
Volzhsko-Kamsky Territory from the Samara bend and Buzuluksky
coniferous forest.
The species is divided into 6 subspecies which are different,
in particular, by the above-described types of the coloration
of the body and the size proportions. E. a. arguta (Pallas, 1773)
– is distributed in the western Kazakhstan. E. a. deserti (Gmelin,
1789) – occupies the whole western part of the distribution range
from the river Ural in the east up to the eastern part of the North
Caucasus in the south-west. E. a. transcaucasica Darevsky, 1953
inhabits eastern Transcaucasia, E. a. uzbekistanica Chernov, 1934
– Uzbekistan and the frontier regions of Turkmenistan, southern
Kazakhstan, Kyrgyzstan and Tajikistan. E. a. darevskii Tsaruk, 1986
– Issyk Kul hollow in Kyrgyzstan. E. a. potanini Bedriaga, 1912 lives
in the area adjacent to the Balkhash Lake and Zaisan hollow in
Kazakhstan.
CONSERVATION STATUS. The steppe – runner E. a. potanini
is included into the Red Data Book of Mongolia (1997) as a
rare species, it is protected on the territory of the Great Gobi
reservation. E. a. uzbekistanica is included into the Red Data Book
of Turkmenistan (1999) as a species which habitats are destroyed
the number of is reducing (category 2). E. a. transcaucasica is
included into the Red Data Book of Armenia (1987) with the status
“species disjuncted from the main distribution range”. E. a. deserti
is included into the Red Data Book of Moldova (2001): category
– Critically Endangered species.

TERRA TYPICA. Karakum, Turkmenistan.
DISTRIBUTION. Middle Asia within the plains of Turkmenistan,
Uzbekistan, Tajikistan, middle and southern Kazakhstan. The
northern border of the distribution range runs in Kazakhstan, in
some places nearly attaining 49°N. Outside of Middle Asia it also
occurs in the north-east of Iran (eastern Khorasan), in the adjacent
plain regions of Afghanistan to the north of Hindu Kush, as well
as in the east up to the western regions of the Xinjiang-Uyghur
Autonomous Region, China.
CONSERVATION STATUS. The number is stable and does not require
special measures on its maintenance.

TERRA TYPICA. Kyzyl Kum, region of Aral-Caspian deserts.
DISTRIBUTION. Middle Asia within the limits of Turkmenistan,
Uzbekistan, extreme west of Tajikistan and Kazakhstan. The northern
border of the distribution range stretches in Kazakhstan from
the north-eastern coast of the Caspian Sea and northern coasts
of the Aral Sea to the north of the Balkhash Lake across the desert
Betpack-Dala up to the basin of the Lake Alakol’. CONSERVATION STATUS. The number is stable. The species does
not require species conservation measures.

TERRA TYPICA. Settlement Sary-Jaz, ravine Terekty, Kyrgyzstan.
DISTRIBUTION. This form of the complex “Eremias multiocellata”
is known from the central Tien Shan (basin of the river Sary-Jaz),
Kyrgyzstan and China (Xinjiang-Uyghur Autonomous Region, including
Kashgaria in the southern Tien Shan). Populations of this
species were referred earlier to Eremias buechneri.
CONSERVATION STATUS. Undetermined.

TERRA TYPICA. Betwen Faizabad and Nuzi, eastern Iran.
DISTRIBUTION. The republics of the Middle Asia within
Turkmenistan, Uzbekistan, southern and south-eastern Tajikistan
and south of Kazakhstan, where it occurs isolatedly in the area
adjacent to the Balkhash Lake. Besides, it is distributed in the
north-eastern Iran (Khorasan) and northern Afghanistan.
CONSERVATION STATUS. The species does not require special conservation
measure.

TERRA TYPICA. Desert Gobi, the road between Sume and river
Tola, Mongolia.
DISTRIBUTION. It inhabits montane and partly submontane regions
of Tien Shan and Pamir-Altai in Kyrgyzstan, eastern and
south-eastern Kazakhstan and neighbouring Uzbekistan. It is
known in Tuva and widely distributed in Mongolia (Bayan Ulug,
Kobdo, Ubsunur, Dzabkhan, Gobi-Altai, Bayan-Khongor, Uver-
Khangai, South-Gobi, Eastern-Gobi and Middle-Gobi aimaks). It is
also known from north-western China southwards to the mountain
ranges of Nan Shan and north-eastern Tibet, in the desert Alashan
and the southern part of Mongolia in the east. During recent years
the taxonomic status of some subspecific forms was revised, they
are considered as separate species.
CONSERVATION STATUS. As a narrow-ranged and small in number
species it is included into the Red Data Book of Kazakhstan (1996):
category 4 – a rare species, known by single specimens.

TERRA TYPICA. Sistan, Iran.
DISTRIBUTION. It occurs in the extreme south-east of Turkmenistan,
in southern Uzbekistan and in the south-west of Tajkistan up to
Kurgan-Tyube and Pyandzh river in the north-east of Iran and in
the north of Afghanistan.
CONSERVATION STATUS. The species is included into the Red Data
Book of Turkmenistan (1999): category 2 – species, the habitats
and number of which are sharply reducing, and into the Red Data
Book of Uzbekistan (2003) – vulnerable, reducing, mosaically
distributed species (2/VU:D).

TERRA TYPICA. Settlement Tengizbai, Altai Range.
DISTRIBUTION. It occurs in the mountains of Tien Shan system and
northern Pamir-Altai within the limits of Kyrgyzstan, Ferganskaya
valley in Uzbekistan and northern Tajikistan. The record of this
species in the extreme south-east of Kazakhstan (Terskei-Alatau)
needs to be confirmed more precisely.
CONSERVATION STATUS. The species does not require special conservation
measures.

TERRA TYPICA. Isfahan city, central Iran.
DISTRIBUTION. It occurs in the south of Turkmenistan where two
isolated populations are known – in the valley of the river Kushka
and the depression Er-Oilan-Duz in the Badkhyz Nature Reserve.
The main distribution is situated within Iran, in the central and eastern
parts of the Iran Plateau. CONSERVATION STATUS. A common species on the territory of
Turkmenistan. Species with a stable number lives on the protected
territory of the Badkhyz Nature Reserve.

TERRA TYPICA. Town Nakhichevan, Azerbaijan.
DISTRIBUTION. It occurs in the left-bank valley of the river Aras in
Armenia and in Nakhichevan, Azerbaijan, i.e. approximately from
the town Echmiadzin in the west up to the southern foothills of the
Megrinsky range. Outside of the eastern Transcaucasia it occurs in
the extreme east of Turkey and in the north-western Iran.
CONSERVATION STATUS. The species does not require special conservation
measures.

TERRA TYPICA. Alashan desert.
DISTRIBUTION. It is distributed mainly in the north China (Inner
Mongolia, Xinjiang-Uyghur Autonomous Region) and in Mongolia,
from where it penetrates into the territory of the neighbouring Tuva.
In North Eurasia the species is widely distributed in the western
and southern Mongolia (Ubsunur, Kobdo, Dzabkhan, Gobi-Altai,
Bayan-Khongor, Uwer-Khangai, South-Gobi, East-Gobi, Middle-
Gobi aimaks) and in Tuva [in Russia]. In the flood-plain of the river
Nariin-Gol lives the subspecies E. p. tuvensis Szczerbak, 1970.
CONSERVATION STATUS. E. p. tuvensis is included into the Red Data
book of the Russian Federation (2001): category 3 – rare species
within the limits of Russia known on the northern periphery of the
distribution range.

TERRA TYPICA. Town Shirabad, Surkhandar’inskaya region,
Uzbekistan.
DISTRIBUTION. It occurs in the extreme south-east of Turkmenistan,
south of Uzbekistan and in the south-western Tajikistan. Outside
of Middle Asia the Tajik Racerunner is known from northern
Afghanistan.
CONSERVATION STATUS. The species is included into the Red Data
Book of Turkmenistan (1999) as a rare species – category 3.

TERRA TYPICA. Region between the Aral and Caspian Seas.
DISTRIBUTION. The species is distributed in sandy deserts
of Middle Asia, Kazakhstan, eastern Iran, adjacent regions of
Afghanistan and northern Belujistan (Pakistan). In North Eurasia it
occurs in Turkmenistan, Uzbekistan, western Tajikistan and southern
Kazakhstan. The northern border of the distribution range
runs in Kazakhstan, from north-western coasts of the Caspian Sea
along the northern area adjacent to the Aral Sea and southern
area adjacent to the Balkhash Lake up to the basin of Alakul Lake
in the east.
Three subspecies are described, which differ mainly by
the color pattern of the dorsal surface of the body. E. s. scripta
Strauch, 1867 occupies the most part of the distribution range
in Turkmenistan and Kazakhstan. E. s. lasdini Tzarevsky, 1918 lives in the south-east of Uzbekistan and south-western Tajikistan.
E. s. pherganensis Szczerbak et Washetko, 1973 is known from
Ferganskaya valley, mainly in the northern Tadzhikistan.
CONSERVATION STATUS. Fergana Sand racerunner E. s. pherganensis
is included into the Red Data Book of Uzbekistan (2003):
category 1, EN – endangered, locally distributed endemic subspecies,
and of Tajikistan (1997) – the category an endangered
subspecies.

TERRA TYPICA. Town Echmiadzin, Armenia.
DISTRIBUTION. In North Eurasia the species is distributed in the
eastern Transcaucasia: the valley of the middle current of Aras river,
from the south-eastern foothills of the Karabakh range and the
valley of Bargushat in the south-east up to the southern spurs of
the mountain Aragaz and Oktemberyanskaya steppe in the west,
in the Karabakh desert and on the plateau Zuvand, as well as in
the south-western Turkmenistan (southern slopes of the mountain
ranges Kyupendag, Karagez and Kopet Dagh). Eremias strauchi
occurs in eastern Turkey and in the north-west and north-east of
Iran, in Iranian Azerbaijan.
The species is differentiated into two subspecies. The nominative
subspecies is distributed in the eastern Transcaucasia, as well
as in the eastern Turkey and Azerbaijani provinces of north-western
Iran. The second subspecies, E. s. kopetdaghica Szczerbak
1972 is distributed in Kopet Dagh, northern Khorasan and eastern
Mazanderan (Turkmenistan and Iran).
CONSERVATION STATUS. The species does not require special protection
measures.

TERRA TYPICA. Kyrgyzstan, region of the Lake Issyk Kul, vicinity of
the town Przhevalsk.
DISTRIBUTION. This form of the complex “Eremias multiocellata”
is distributed in the basin of the Lake Issyk Kul.
CONSERVATION STATUS. The species does not require special protection
measures.

TERRA TYPICA. Kyrgyzstan, valley of Arpa River.
DISTRIBUTION. This form of the complex “Eremias multiocellata” is
distributed in the basin of the river Naryn in Kyrgyzstan, the valley
of the rivers At-Bashi and Arpa. Foothills of Moldo-Too, Baibishe-
Too, Yaman-Too, At-Bashi and Naryn-Too.
CONSERVATION STATUS. Not determined.

the Ural River).
DISTRIBUTION. Rapid racerunner is a widely distributed species,
occurring from the eastern part of the North Caucasus and lower
region of the Volga River to northern Iran, Afghanistan and eastwards
up to north-western China (Xinjiang-Uyghur Autonomous
Region., Inner Mongolia and Hansu Province). In North Eurasia
the distribution range of the species stretches into the eastern
Ciscaucasia, lower region of Volga River, eastern Transcaucasia in
the lowland Georgia and Azerbaijan, Middle Asia and Kazakhstan.
Several subspecies are described, of which three subspecies occur within the limits of the countries of North Eurasia. E. v. caucasica
Lantz, 1828 inhabits eastern Ciscaucasia and Transcaucasia.
The most part of the distribution range is occupied by the nominative
subspecies E. v. velox, and in the Issyk Kul hollow occurs the
subspecies E. v. borkini Eremchenko et Panfilov, 1999, described
from the Tien Shan mountains, in Kyrgyzstan (southern shore of
the Lake Issyk Kul, from the region between the rivers Ak-Terek
and Ak Sai).
CONSERVATION STATUS. The species does not require special protection
measures.

TERRA TYPICA. Eastern Turkestan, Yarkend, Kashgaria, China
(Xinjiang-Uyghur Autonomous Region).
DISTRIBUTION. The main distribution range of the species is
situated in north-western China (Xinjiang-Uyghur Autonomous
Region, Inner Mongolia) and in Mongolia (Gobi-Altai, Bayan-
Khongor, South-Gobi aimaks). An isolated population lives in the
Zaissan hollow in the south-east of Kazakhstan.
CONSERVATION STATUS. The species is included into the Red Data
Book of Kazakhstan (1996): category 4 – not investigated species
with a limited distribution range.

TERRA TYPICA. Town Yarkend, Kashgaria, China (Xinjiang-Uyghur
Autonomous Region).
DISTRIBUTION. The distribution range of this species is marked by
the Tarim basin which includes the southern slopes of Tien Shan,
piedmont plains and foothills of Kunlun and eastern Pamir (excluding
sands Takla-Makan) in China and eastern Alai in Kirghizia. Earlier
this form was considered as a subspecies of multiocellated racerunner
E. multiocellata Gunther, 1872.
CONSERVATION STATUS. Not determined.

TERRA TYPICA. South Sweden.
DISTRIBUTION. Sand lizard is one of the most widely distributed
Eurasian species; its distribution range covers the most part of
Europe from western France and north of the Balkan Peninsula to
Eastern Siberia, north-western Mongolia and western China in the
east (Chinese part of the Mongolian Altai). On the territory of North
Eurasia the species occurs from western borders of Moldova, the
Ukraine, Belarus, the Baltic States and Russia in the west to the
northern area adjacent to Lake Baikal and southern Trans-Baikal area
in the east, more or less in parallel with the north border of taiga. It
inhabits the Caucasus and Kazakhstan in the south. It is known from
Mongolia (Kobdo aimak). In the south-east distribution range covers
the eastern Semirechye, north-western Mongolia (Kobdosky
aimak) and the western part of Xinjiang-Uyghur Autonomous
Region in China. From there it penetrates into Issyk-Kul depression
in Kyrgyzstan. Data on the records in piedmonds of Turkestan and
Seravshan mountain ranges in eastern Uzbekistan are erroneous
Ten subspecific forms are distinguished, the following seven
occur in the former Soviet Union. L. a. chersonensis Andrzejowsky,
1832 – Moldova, the right-bank Ukraine, Belarus, the Baltic States,
Leningrad region and south of the neighbouring Karelia. In the
east, approximately from the left-bank valley of the river Dnieper
a narrow area of intergradation with the neighbouring eastern
subspecies is noted. L. a. exigua Eichwald, 1831 occupies the
whole eastern part of the distribution range up to the Crimean
Peninsula and Ciscaucasia in the south. L. a. grusinica Peters, 1960
inhabits the coast of the Black Sea and submontane regions of the
Caucasus in the south-west of the Krasnodar Territory, Abkhazia,
in the Colchis lowland and Ajaria. L. a. brevicaudata Peters, 1958
occurs in northern and western Armenia, southern Georgia and
on the southern slopes of the Great Caucasus range within the
North Ossetia. L. a. ioriensis Peters et Muskhelischwili, 1968 occurs
on the southern slopes of the Caucasian range: in the valley and
ravine of the upper current of the river Iori in Georgia. L. a. boemica
Suchow, 1929 inhabits submontane regions of North Ossetia,
Ingooshetia, Chechnya and Dagestan. L. a. tauridica Suchow, 1926
inhabits the southern montane Crimea.
CONSERVATION STATUS. The species is included into the Red
Data Book of Latvia (2003) and Estonia (1998). Its conservation
status (category 2) corresponds to the same status of the Bern
convention.

TERRA TYPICA. Tiflis city (=Tbilisi), Georgia.
DISTRIBUTION. The most part of the distribution range in
North Eurasia is situated in the Caucasus. There it is noted in the
Transcaucasia, in eastern and northern Georgia, submontane and
montane regions of Azerbaijan and Armenia, as well as in river
valleys in interior Dagestan. Besides, it occurs on the coast of the
Black Sea of the Krasnodar Territory, Abkhazia and Ajaria. Outside
of the Caucasus it is encountered in the eastern Turkey and northwestern
Iran. One of the three subspecies, L. m. media Lantz et
Cyren, 1920, occurs in the Caucasus.
CONSERVATION STATUS. The species is included into the Red Data
Book of the Russian Federation (2001): category 3 – a rare species
within the limits of Russia with distribution on the extreme northern
border of its range.
Female (left) and male (right) of Medium Lizard

TERRA TYPICA. Krasnovodsk, southern Turkmenistan.
DISTRIBUTION. L. strigata is widely distributed in North Eurasia. It
lives predominantly in the eastern half of the Caucasus, at least up
to the latitude of the towns Armavir and Stavropol in the Stavropol
Territory in the north-west and the valley of the river Terek and
lower reaches of the river Kuma in Dagestan in the north-east. In
the eastern Transcaucasia it mainly occurs in the lowland regions
of Azerbaijan, Armenia and Georgia, where it is not known to the
west of the Suramsky range. Until recently an isolated population
existed in the area of the cape Pizunda in Abkhazia. It also occurs
in the south-west of Turkmenistan. Outside of the Caucasus it is
known from eastern Turkey and northern Iran.
CONSERVATION STATUS. On the most part of the distribution range
the number of the species is stable.

TERRA TYPICA. Vienna, Austria.
DISTRIBUTION. It is distributed in middle and southern Europe, in
the north-western part of Asia Minor, on the most part of Moldova
and south-west of the right-bank Ukraine. On the valley of the river
Dnieper it nearly reaches Kiev. In its middle current it appears in
the left-bank area, where on the valley of the river Vorskla reaches
the Poltava city in the region with the same name. Of five described
subspecies L. v. viridis Laurenti, 1768 occurs in the Ukraine
and in Moldova.
CONSERVATION STATUS. On the most part of the distribution range
the number of the species is stable.

TERRA TYPICA. Basminsk, between Tebriz and Teheran, northern
Iran.
DISTRIBUTION. The species is distributed in the north-western
Iran, in the provinces Eastern Azerbaijan and Esfahan. On the territory
of North Eurasia the only record is known in 1880 in the
south-eastern Azerbaijan (Talysh).
CONSERVATION STATUS. Uncertain due to poor knowledge on the
species.

TERRA TYPICA. Kaiseri, Asia Minor (Turkey).
DISTRIBUTION. It is widely distributed in Asia Minor, from where
it also penetrates into central and northern Armenia. At present
the species is preserved mainly in the Spitak region where it is observed
only on areas of difficult access and on unplowed areas.
CONSERVATION STATUS. It is included into the Red Data Books of
the USSR (1984) and Armenia (1987) as an endangered species. It
is protected by the Bern Convention. This species was referred to subgenus Parvilacerta Harris,
Arnold et Thomas, 1998. This subgenus includes 2 species:
Parvilacerta fraasii (Lehrs, 1910) and Parvilacerta parva
(Boulenger, 1887).

TERRA TYPICA. Synd, between Karachi and Sukkur.
DISTRIBUTION. The species is widely distributed on the
whole Iranian Plateau and in Afghanistan, comes into southern
Turkmenistan. In North Eurasia it occurs only in the south of
Turkmenistan up to the vicinity of the town Chärjew in the east.
Previously this form was considered as a subspecies of M. guttulata.
CONSERVATION STATUS. In Turkmenistan the status of the populations
is rather stable and does not require special conservation
measures. It is protected on the territory of the Badkhyz Nature
Reserve.

TERRA TYPICA. Vicinity of Baku city, Azerbaijan.
DISTRIBUTION. The species is widely distributed in the northeast
of the Balkan Peninsula, some islands of the Aegean and
Mediterranean Seas, Sinai Peninsula, Asia Minor, Middle East
and the Caucasus to Pakistan and north-western India in the
east. On the territory of North Eurasia it occurs in the Caucasus
in Azerbaijan, Armenia and eastern Georgia, approximately to
the city Tbilisi in the west. There are data which require confirmation
about the occurrence of an isolated population in the North
Caucasus in the submontane Chechnya. Of six known subspecies
only O. e .elegans occurs in the Transcaucasia.
CONSERVATION STATUS. The status of the populations on the
whole distribution range is stable and does not require special
protection measures.

TERRA TYPICA. North of the Crimean Peninsula.
DISTRIBUTION. The distribution range of the species covers the
countries of the Balkan Peninsula, western Turkey, as well as the
south of the Ukraine and Moldova. It is widely distributed in the
Crimea, in the Kherson, Nikolaev and Odessa regions in the southwest
of the Ukraine and in the south of Moldova. Of several known
subspecies P. t. taurica occurs in the Ukraine and in Moldova.
CONSERVATION STATUS. The species does not require special
measures on its protection.

TERRA TYPICA. Amur region.
DISTRIBUTION. It occurs in north-eastern China, Korea and Japan
(the island Zussima). In North Eurasia it lives in Russian Far East – in
the southern regions of the Primorsky and Khabarovsk Territories.
CONSERVATION STATUS. The species does not require special
measures on its protection.

TERRA TYPICA. Chemul’po, Korea.
DISTRIBUTION. It occurs in Korea, south-east Manchuria and eastern
China from where it penetrates into the south of the Primorsky
Territory approximately to the valley of the river Iman in Russia.
CONSERVATION STATUS. Not determined.

Vienna, Austria.
DISTRIBUTION. One of the most widely distributed Eurasian
species. A common species in the northern half of Eurasia from
Ireland and the Pyrenean Peninsula in the west to the Shantarskie
islands, the Sakhalin island and northern Japan in the east. It occurs
in Mongolia, in the Mongolian Altai, in Bain-Ulug, Khubsugul
and Central aimaks, China (Chinese Altai in the Xinjiang-Uyghur
Autonomous Region). In Russia the northern border of the distribution
range from the coast of the Kola Peninsula in the north-west
continues behind the Polar Circle up to the lower current of Yenisei
River. Further to the east it crossed the valleys of Lena River and its
tributaries (Vilyui and Aldan). In Far East it comes out to the sea
somewhat to the south of the valley of the river Uda. The southern
border of the distribution range from the Trans-Carpathians
continues to the east between forest-steppe and steppe. In
Ukraine the most southern, evidently isolated habitats are known
in the Novomoskovsk district of the Dnepropetrovsk region. It occurs
everywhere on the Sakhalin Island. On the whole extensive
distribution area of the viviparous lizard four subspecies are distinguished:
Zootoca vivipara carniolica Mayer, Böhme, Tiedemann
& Bischoff, 2000, Zootoca vivipara pannonica (Lac & Kluch, 1968),
Zootoca vivipara sachalinensis (Pereleshin & Terentjev, 1963) and
Zootoca vivipara vivipara (Jacquin, 1787). In North Eurasia the
subspecies Z. v. vivipara (Jacquin, 1787) and Zootoca vivipara sachalinensis
(Pereleshin & Terentjev, 1963) are distributed.
CONSERVATION STATUS. The species does not require special
measures on its protection.

TERRA TYPICA. Settlement Nyuvadi in the ravine of the Aras River,
south-east Armenia.
DISTRIBUTION. It is widely distributed in the mountains of Lesser
Caucasus, in the most part of the southern and northern Armenia,
southern and southern-east Azerbaijan and the ravine of the upper
current of Kura River in western Georgia. In the Transcaucasia
the distribution range consists of many isolated populations of different
size associated with the ravines of small rivers flowing down
to the river Aras in the south, and tributaries of the river Kura in
the north. Outside the Caucasus it occurs in eastern Turkey (to
the east of the Lake Van) and in the adjacent north-western Iran.
AZERBAIJAN LIZARD DAREVSKIA RADDEI (BOETTGER, 1892)
MEADOW LIZARD DAREVSKIA PRATICOLA (EVERSMANN, 1834)
RI V ER KURA LIZARD DAREVSKIA PORTSCHINSKI I (KESSLER, 1678)
88 88L A
C
E
R
T
I
D
A
E
The southern border of its distribution range is not exactly determined.
Intraspecific taxonomy is still not clear. Three subspecies
are described. The nominative subspecies D. r. raddei inhabits
the most part of Armenia and Azerbaijan, in the basin of the Lake
Sevan in Armenia (mainly in the western part of the republic). In
the ravine of the upper current of the river Kura in Georgia occurs
D. r. nairensis, considered by some taxonomists as a distinct species.
D. r. vanensis (Eiselt, Schmidtler et Darevsky, 1993) occurs
in the eastern part of the basin of Lake Van in Turkey and in the
neighbouring Iran.
CONSERVATION STATUS. In the most part of the distribution range
the number of the species is high and stable.

We tested intra-population morphological variability and the ontogeny of sexual dimorphism in a sample of 78 individuals of Lacerta agilis (Linnaeus, 1758) from the Bistra Mountain in the Former Yugoslav Republic of Macedonia. The effect of predation was also assessed by examining the animals’ tails. In 43.8% of individuals of all age classes tails were missing or regenerated, which means that lizards are subjected to high predation at this locality. After measuring, marking and photographing, all animals were released into their natural habitat. The set of 17 linear measurements and body mass were taken for each individual, and snout-vent length was taken as a standard descriptor of body size. Among adults, significant sexual dimorphism was recorded in body size, head dimensions and distance between front and hind limbs, while sexual dimorphism was not observed in juveniles. Females had greater body size and distance between front and hind limbs, while males had proportionately larger heads. Static allometric trajectories of adult females and males were assessed, and they diverged only for head and mouth width. Divergence of ontogenetic allometric trajectories was significant for all measurements except the dimensions of the fourth toe on the front and hind limbs. Sexual dimorphism pattern in sand lizards from Bistra Mountain was in concordance with previous studies. Larger adult female body size could be interpreted as a result of sexual selection for fecundity, while larger male heads could result from selection for greater bite force in two ways: intra-sexual selection (between males) in relation to territory defense, and inter- sexual in relation to copulatory bites. Sexual dimorphism was not significant among juveniles: adults apparently attain a different morphology through diverging allometric trajectories during growth. Further studies should include populations of sand lizards from different habitat types to further assess patterns of intra-specific morphological variability.

To investigate the impact of nuptial coloration (the
badge) on male fitness in the Swedish sand lizard
(Lacerta agilis), we conducted a manipulation
experiment in a natural population. Males in one
group had their badges enlarged by being painted as
cheaters and were compared to a control group with
respect to mate acquisition, body condition and survival.
Badge enlargement did not affect survival, but
elevated mate acquisition by almost 400%, and body
condition in small males. This increase in condition
is likely to stem from greater access to female-associated
sites with high food availability.

One hundred seventy-seven species of amphibians and reptiles in sixty-fiv genera have been recorded from within the political boundaries of Iran. The actual number of species may prove to be somewhat less when problems of synonymy have been more extensively investigated. Thirty-three species or 19 per cent of the known herpetofauna, are represented in the present collection. The majority of these (21 species) were collected in the western foothills of the Zagros Mountains in southwestern Iran.
Relatively few collections have been made in Iran, and many species have been recorded from single localities. Many taxonomic problems must await the collection of further specimens and examination of typical material. Distribution is incompletely known for all species. Many of the early locality records are not precise, and require confirmation.
The aridity and prolonged hot season impose severe limitations on the fauna of the foothill region of southwestern Iran. The length of the growing seasonn varies with the duration of the winter and spring rainy season. The vegetation is almost entirely herbaceous in this area, and is extensively grazed by sheep and goats. By late spring the hills are completely denuded, The large numbers of insects, which coincide with the blooming of herbaceous plants in the spring, dwindle through the summer. Grasshoppers and ants form the bulk of the food supply for many species throughout the summer.
Various behavioral adaptations enable the amphibians and reptiles to survive the long summer. Foraging activity begins in the early morning hours, diurnal lizards quickly reaching temperatures of normal activity by exposing a maximum surface area to the direct rays of the sun. As ambient temperatures rise, the period of activity is extended by utilization of small areas of shade (particularly the case with lacertids), brief dashes being made into direct sunlight. The agamid lizards may remain in the sun, but position themselves so that the sun´s rays strike a minimum surface area, and at a small angle. Agama agilis ascends low bushes during extremely hot periods, and thus may escape much of the reradiated heat from the hot ground surface. The agamid lizards may benefit from changes in their albedo. They are dark in color when basking, and much lighter during the heat of the day. These color changes may supplement behavioral temperature regulation.
The reptiles retreat to shelter during the hottest hours. Deep crevices in limestone outcrops provide shelter for Agama nupta, Mabuya aurata septemtaeniata, and several of the snakes. Agama agilis and Eremias guttulata watsonana seek refuge under small rocks or in rock piles on level ground. The latter species also utilizes small holes and cracks in the hard-packed soil.
A few small streams persist throughout the summer, ofter only as intermittent pools. In many dry stream beds, moisture may be found a few inches beneath the surface. A moist, cool, microclimate is also maintained in small caves and ssinkholes in the gypsium formations. These areas provide a summer refuge for the amphibians, and shelter nocturnal reptiles during the day. These cracks and crevices are also utilized by Agama nupta during the hottest hours, and presumably at night.
Several species of geckos and snakes are able to exist in the area due to their nocturnal habits, and the fact that temperatures do not drop below their normal activity range at night during the summer.
Dune-dwelling species are able to escape the severe midday temperatures by retreating the burrows a few inches below the sand surface.
Anal temperatures indicate that the normal activity range for most of the diurnal lizards of southwestern Iran is between 38.0° and 42.0°C. Uromastix loricatus was actively foraging at times when its anal temperature exceeded 43°C. Agama agilis was consistently found in direct sunlicht, the anal temperature above 40°C. Critical maximum temperatures for most diurnal lizards are between 46° and 49°C. Both Uromastix loricatus and Scincus conirostris recovered from temperatures above 50°C. Cowles and Bogert (1944) found that the normal activity range of diurnal lizards of the deserts of the southwestern U.S. was approximately 35°-43°C., and that the lizards preferred temperatures of about 37°C. They found that lizards avoided temperatures above 40°C. Critical maximum temperatures were reported to bte 43°-49°C.
As air and surface temperatures increased during the late spring and summer in southwestern Iran, a shift in the periods of reptilian activity was noted. During the moderate spring temperatures, activity was confined to the midday hours. By midsummer, foraging activity was noted only during the earliest daylight hours,a nd again in late afternoon.
A detailed zoogeographic analysis of the Iranian herpetofauna must await more extensive locality data, and the analysis of many systematic problems. Nevertheless, it is apparent that the several geographic and ecologic provinces of Iran are characterized by distinct differences in the composition of their faunas. Southwestern Iran and the northwestern highlands have a greater number of species with Mediterranean affinities than do the regions east of the Zagros Mountains. Southeastern Iran shares species with Pakistan and India, while elements of the Central Asia fauna are present in the northeast of the country. Apparently there is a relatively high degree of ebndemism on the central plateau of Iran. A European fauna prevails north of the Elburz Mountains on the south coast of the Caspian Sea. The majority of amphibians and reptiles known Iran are restricted in their distribution to the Middle East.
Southwest Asia has apparently existed as a geographic unit long enough to have developed a characteristic herpetofauna, particularly at the species level.
Acanthodactylus cantoris blanfordi, Acanthodactylus cantoris schmidti, Eremias guttulata watsonana, Ophisops elegans elegans.

Anderson, S.C. (1968) -

Zoogeographic analysis of the lizard fauna of Iran. - In: Fisher, W.B. (ed.) The Cambridge history of Iran, vol. I, The land of Iran. Cambridge University Press, London: 305-371.

×

Anderson, S.C. (1968) -

Chapter 10: Zoogeographic analysis of the lizard fauna of Iran. - In: The Cambridge History of Iran, Volume I: The Land of Iran. pp. 305-371.

The collection of amphibians and reptiles made by the Street Expedition to Afghanistan 1965 consists of 247 specimens, including six species of amphibians and 38 species of reptiles. Definite Afghan localities are recorded for the first time for the following species: Agama agrorensis, Agama erythrogaster, Agama himalayana, Agama lehmanni, Agama nuristanica (new species herein described), Agama abdakhshana (new species herein described), Cyrtodactylus watsoni, Hemidactylus flaviviridis, Eremias nigrocellata, Eryx elegans (type was recorded as coming from Afghanistan, but n precise locality was given, An analysis of the zoogeographic significance of this and other collections from Afghanistan is contemplated for the future.
Acanthodactylus cantoris, Eremias guttulata watsonana, Eremias nigrocellata, Eremias velox persica.

Sex steroids levels in the plasma and testis during the reproductive cycle of lizard Podarcis s. sicula Raf. - General and Comparative Endocrinology, 85 (1): 1-7.

×

Progesterone (P), 17-OH-progesterone (17-OH-P), androstenedione (A), dehydroepiandrosterone (DHEA), testosterone (T), 5α-dihydrotestosterone (5α-DHT), and 17β-estradiol (E2) were measured by RIA in plasma and testes of 114 males of the oviparous lizard Podarcis s. sicula raf, a species that displays annual hibernating cycles. Hormones were determined each month from January until December, except for August. Testosterone peaked at 174.8 ng/ml of plasma after emergence (March), while 5α-DHT and A peaked in April. Plasma DHEA increased during hibernation. During the refractory period there were progressive increases in P and E2 plasma levels. The testicular peak of T, in March, coincided with that observed in plasma. The striking increases in testicular T and A in early July occurred at a time when plasma androgen concentrations were low. 5α-DHT increased in April when spermatogenesis with spermiation occurred and then decreased alongside a second peak of T. There is an apparent separation of plasma and testicular androgen concentrations during the reproductive cycle.

Progesterone, 17-hydroxyprogesterone, androstenedione, 5 alpha-dihydrotestosterone, dehydroepiandrosterone, testosterone and oestradiol concentrations in the plasma were measured by simultaneous radioimmunoassay in males of the lizard Podarcis s. sicula. Hormonal determinations were performed at monthly intervals from January to December (except for August). Testosterone and androstenedione reached peak values of 174.8 ng/ml and 21.4 ng/ml in the mating season (spring) and then testosterone fell abruptly to 5.9 ng/ml in June remaining at this level during hibernation when dehydroepiandrosterone (DHA) reached a maximal level of 28.5 +/- 9.3 ng/ml. Castration resulted in a marked decrease of testosterone, androstenedione, dihydrotestosterone and DHA values, with DHA being significantly lowered only during the winter season. In castrated animals, however, testosterone and androstenedione persisted conspicuously in the plasma during the breeding period, suggesting that adrenal sex steroid output may change during the annual reproductive cycle. In intact animals, progesterone and oestradiol exhibited peak values during the refractory period after the mating season. We suggest a probable role of oestradiol in the induction of the refractory period in this lizard.

The author updates the knowledge on the herpetofauna of southern Dobruja according to his
own studies within the area, during 1993-1998, when he identified 18 taxa. They represent 37.5% of
the total amphibian and reptile species from Romania. Also the author makes some remarks on
respective ecosystems and on the urgent protection need of some species.

Andrén, C. (2004) -

Oskarshamn site investigation Amphibians and reptiles in SKB special area of investigation at Simpevarp. - Svensk Kärnbränslehantering AB. P-04-36. 30 pp.

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Amphibian and reptile species likely to occur in the Simpevarp SKB special
area of investigation are listed with comments on their distribution, status, biology
and environmental demands. The species are Triturus vulgaris (Smooth newt),
T cristatus (Great crested newt), Bufo bufo (Common toad), Rana arvalis (Moor frog),
R temporaria (Common frog), Lacerta agilis (Sand lizard), L vivipara (Common
lizard), Anguis fragilis (Slow-worm), Coronella austriaca (Smooth snake), Natrix
natrix (Grass snake) and Vipera berus (Adder). A short field study was performed
mainly to verify the presence of suitable habitats for the species listed. Findings of
amphibians and reptiles as well as their potential habitats are noted by geographical
codes and additional remarks are given to interesting findings. Altogether 38 findings
of five different species of amphibians and reptiles were done in 18 localities. Of special
interest is the record of sand lizard, which probably indicate a colony in the vicinity
even though this could not be confirmed.

The Sand lizard Lacerta agilis inhabits a huge area across Eurasia with several subspecies. Nine subspecies are generally approved, L. a. agilis, L. a. argus, L. a. bosnica; L. a. chersonensis, L. a. exigua, L. a. grusinica, L. a. iorinensis, L. a. brevicaudata and L. a. boemica, but several more have been described. The emergence of this large number of subspecies is connected with the phylogeographic history of this species, defined by geographic and climatic processes. A study revealing phylogenetic relationships of this species was already conducted with a broad taxon sampling and coverage. However, the latter study was based solely on the cytochrome b gene and, furthermore, the Balkan Peninsula inhabited by the subspecies L. a. bosnica was underrepresented. This peninsula is a centre of European herpetofaunal endemism with high levels of phenotypic and genotypic variation. Therefore, the inclusion of the subspecies L. a. bosnica is important to clarify the overall view of the phylogenetic relations within the Lacerta agilis subspecies complex and to investigate the level of population differentiation within this highly diverse area. Thus, the aim of the present study was a more thorough analysis of the Balkan Peninsula with a broader taxon sampling. Furthermore, we extended the available datasets by adding the mitochondrial d-loop region and by further samples of different areas of the distribution range apart from the Balkan Peninsula. Our study reveals that the Balkan Peninsula is apparently inhabited by several differentiated lineages, whereby the Central Greek lineage might deserve subspecies status. Furthermore, the distribution area of the two subspecies L. a. agilis and L. a. argus should be revised, as the results of our study raise doubt about the commonly assumed distribution of both subspecies. As the most important outcome our results support that L. a. boemica deserves species status.

The sand lizard (Lacerta agilis) inhabits a huge area across Eurasia with several subspecies. Nine subspecies are generally approved, L. a. agilis, L. a. argus, L. a. bosnica; L. a. chersonensis, L. a. exigua, L. a. grusinica, L. a. iorinensis, L. a. brevicaudata and L. a. boemica, but several more have been described. The emergence of this large number of subspecies is connected with the phylogeographic history of this species, defined by geographic and climatic processes. Within Germany two subspecies exist according to Bischoff (1988), L. a. argus und L. a. agilis, which are separated based on morphological features. However, the existence of these two subspecies was questioned by Rahmel (1988) and by Elbing et al. (1996). We conducted phylogenetic analyses based on the cytochrome b gene sequences from specimen of L. a. agilis and L. a. argus with samples covering their whole distribution range. These sequences already have been published before by Andres et al. (2014) and Kalyabina-Hauf & Ananjeva (2004). Our study challenges the commonly assumed distribution of L. a. agilis und L. a. argusand their status as distinct subspecies. Therefore their genetic status and distribution range should be thoroughly analysed and revised in future studies.

In contrast to the view of a diffuse monoaminergic innervation of the telencephalon, studies on the monoaminergic innervation in certain mammalian isocortical regions have shown a high degree of regional and laminar specificy. The present study was designed to examine the distribution patterns of dopamine, noradrenaline and serotonin in a telencephalic structure, the anterior dorsal ventricular ridge, of the sand lizard Psammodromus algirus (Lacertidae) using specific antibodies against each monoamine.
The anterior dorsal ventricular ridge receives an abundant monoaminergic innervation compared to that of cortical telencephalic regions. The distribution of the different monoamines presented zonal and regional patterns throughout the ridge. The cell cluster zone was profusely innervated by catecholamines, whereas no serotoninergic fibers innervated the cell bodies in the cluster zone. On the other hand, the periventricular zone was heavily innervated by serotonin, but catecholaminergic fibers were almost lacking. With regard to regional patterns, dopamine exhibited major differences in the mediolateral axis of the anterior dorsal ventricular ridge: dopaminergic innervation was densest in the lateral region, which in other reptiles is described as a target of visual thalamic projections. Whereas the zonal pattern of the monoaminergic innervation of the anterior dorsal ventricular ridge seems to be a constant feature in the reptiles studies to date, the regional pattern varies among reptilian groups, especially taking into account the density of monoaminergic innervation.

We have studied the intrinsic connections of the anterior dorsal ventricular ridge (ADVR) in the lacertid lizard Psammodromus algirus by means of retrograde transport of horseradish peroxidase (HRP) and fluorescent labeling with the lipophilic carbocyanine dye DiI. We injected HRP into different regions in the ADVR arrayed in a medial-to-lateral sequence, with each consisting of three distinct superficial-to-deep zones. When HRP was injected into a given region, many labeled neurons (always located ipsilateral to the injection site) were found at all mediolateral regions of ADVR in locations rostrally distant from the injection site. DiI crystals were applied on different superficial-to-deep zones within each region. Two patterns could be recognized: DiI crystals applied on the periventricular (most superficial) zone resulted in a labeling of cells widely distributed throughout the ADVR independently of the mediolateral region of the application site, whereas DiI crystals applied on deeper zones resulted in a staining of cells mostly restricted to a narrow radial area. Results from both types of labeling confirm that the ADVR has a prominent radial component in its intrinsic organization, but they also demonstrate that some areas of the ADVR receive projections from distant, rostrally located neurons in every ipsilateral region of the ridge itself, which establishes a clear non-radial component. This organization may have important functional properties with regard to a putative integration of different sensory modalities conveyed by thalamic afferent fibers to the ADVR. Last, we analyzed some evolutionary implications of our results.

Ultrastructural studies suggest that, in the oocytes of the lizardPodarcis sicula, ribosomal bodies are structurally continuous with annulate lamellae during their organization and disaggregation. This observation may indicate the dynamic transformation of the cytomembranes of one structure into those of the other, and vice versa. Moreover, the presence of annulate lamellae has been detected for the first time in lizard oocytes. The hypothesis is advanced that ribosomal bodies and annulate lamellae, in spite of some different structural characteristics, may play a similar role during the oocyte growth.

Role of phylogenetic relations and conditions of habitat of lizards of „Lacerta saxicola“ complex in formation of the heat resistance level of tissue as specific character. - Russian Journal of Herpetology, Moscow, 1 (1): 53-59.

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The extent of resistance of animal cells and tissues to injurious heat effect is a cytophysiological criterion of species. In poikilotherms it can be determined both by phylogenetic relations and ecological conditions of a species` formation and existence. To analyze the relative role of these factors in formation of the tissue heat resistance level as characteristic of a species, a study was made on 9 species of lizards of the `Lacerta saxicola` complex. A characteristic feature of the complex is that some of the species belonging it are of hybrid origin, which implies that genetically they are closely related to other, parental forms. Heat resistance level of muscle tissue in a hybrid species may differ from the parental forms if the conditions of its existence are different. On other hand, genetically unrelated species living under similar conditions may have the same level of heat resistance of muscle tissue. Interspecific differences in heat resistance of muscles cannot be explained by phylogenetic relations only. Of major importance in formation of the cellular heat resistance level in the process of speciation are ecological, and primarily, thermal conditions of the environment.

Abstract
Food habits of the green lizard (Lacerta bilineata) were studied in some Mediterranean sites of the vicinities of Rome (Latium, central Italy), by means both of stomach dissection of individuals found already dead in the field and faeces analysis of living individuals. The taxonomic diet composition of the lizards was accurately described by both methods, thus demonstrating the reliability of faecal pellet analysis as an non-cruel method to study lizard diets. However, by using the faeces analysis technique, it is difficult to measure the exact food intake rate (items / unit of time) as well as the size distribution of prey ingested by lizards. There was a remarkable ontogenetic change in the taxonomic diet composition of green lizards: adults essentially consumed beetles and isopods, whereas juveniles fed mainly upon orthopterans, spiders, and Rhynchota. Cannibalism and predation upon small vertebrates (lizards) were very rare events. It is suggested that (i) age-related differences in microhabitat frequented could explain the dietary differences between young and adult lizards due to different availability of the various prey categories in the various microhabitats, and that (ii) adults and juveniles partition their habitats to reduce intraspecific interference competition.

The evolution of sex steroids in the plasma and gonads of the lizard Podarcis s. sicula during the postreproductive period shows that these hormones could be involved in determining refractoriness. In the male, during this phase, the oestradiol could lower the hypothalamo-hypophyseal system through a negative feed-back. In the female a similar role could be assigned to progesterone.

The effects of mammalian gonadotropins on spermatogenesis in Lacerta s. sicula Raf. was studied during the autumn thermic refractory period.
During this period the mammalian gonadotropins induce a stimulation of spermatogenesis and of the epididymis. FSH acts preferentially on the spermatogonial multiplication and on the primary spermocytes; LH acts on all the stages of spermatogenesis, but in particular on spermiohistogenesis and spermiation; moreover it causes a marked development of the epididymis and of the SSC.
The effects of gonadotrophins are temperature dependent: below 18°C stimulation concerns only the early stages of spermatogenesis and spermiation does not occur; at 27°C a complete spermatogenesis with maximum spermiation occurs. At 30°C gonadotrophins induce a strong stimulation of the gonad, but this stimulation is accompanied by frequent degenerative phenomena of the secondary spermatocytes and spermatids. LH is more capable than FSH of limiting this degeneration. Testosterone does not prevent the regression of spermatogenesis due to high temperatures, while it causes a clear development of the epididymis.

Coloring and drawing of a Sand Lizard (Lacerta agilis) in the territory of the Murom Reserve, Vladimir Region. - In: Specially Protected Natural Territories: Current Status and Development Prospects. Materials of the All-Russian Jubilee Scientific and Practical Conference, dedicated to the 25th anniversary of the Meshchera National Park (October 5-6, 2017). Vladimir: Kaleidoscope, 2018: 85-89.

Previously known in Portugal only in Serra da Estrela (Centre of Portugal), the Iberian rock-li zard, Lacerta monticola, is now reported inside the Montesinho Natural Park (North of Portugal). The specimens found in Museu Bocage`s collection could be tentatively assigned to subspecies call1abrica.

Antunes, P. & Vicente, L. (2001) -

The Museu Bocage new lacertid lizards collection – Established after the 1978 fire. - In: Pérez-Mellado, V. (ed.): Abstracts of the 4th International Symposium on the Lacertids of the Mediterranean Basin, 7-11 May 2001, Mahon, Menorca, Spain: 43.

The diet of both reptiles was studied in the park of the National Agronomic Institute in Alger city. Faeces of the captured animals were analysed in order to evaluate relative abundance of preys, standardised diversity and diet overlaps for both species. They both preyed on a large diversity of small invertebrates and showed overlapping diets. In spite of this opportunistic behaviour, both predator species seem to split the feeding resources of their environment through spatial and temporal segregation.

Geographical body size variation has long interested evolutionary biologists, and a range of mechanisms have been proposed to explain the observed patterns. It is considered to be more puzzling in ectotherms than in endotherms, and integrative approaches are necessary for testing non-exclusive alternative mechanisms. Using lacertid lizards as a model, we adopted an integrative approach, testing different hypotheses for both sexes while incorporating temporal, spatial, and phylogenetic autocorrelation at the individual level. We used data on the Spanish Sand Racer species group from a field survey to disentangle different sources of body size variation through environmental and individual genetic data, while accounting for temporal and spatial autocorrelation. A variation partitioning method was applied to separate independent and shared components of ecology and phylogeny, and estimated their significance. Then, we fed-back our models by controlling for relevant independent components. The pattern was consistent with the geographical Bergmann`s cline and the experimental temperature-size rule: adults were larger at lower temperatures (and/or higher elevations). This result was confirmed with additional multi-year independent data-set derived from the literature. Variation partitioning showed no sex differences in phylogenetic inertia but showed sex differences in the independent component of ecology; primarily due to growth differences. Interestingly, only after controlling for independent components did primary productivity also emerge as an important predictor explaining size variation in both sexes. This study highlights the importance of integrating individual-based genetic information, relevant ecological parameters, and temporal and spatial autocorrelation in sex-specific models to detect potentially important hidden effects. Our individual-based approach devoted to extract and control for independent components was useful to reveal hidden effects linked with alternative non-exclusive hypothesis, such as those of primary productivity. Also, including measurement date allowed disentangling and controlling for short-term temporal autocorrelation reflecting sex-specific growth plasticity.

The effects of immigration on the behaviour of residents may have important implications for the local population characteristics. A manipulative laboratory experiment with yearlings of the common lizard (Lacerta vivipara) was performed to test whether the introduction of dispersing or philopatric individuals influences the shortterm spacing behaviour of resident individuals. Staged encounters were carried out to induce interactions within dyads. The home cage of each responding individual was connected by a corridor to an unfamiliar “arrival cage” to measure the latency to leave their own home cage after each encounter. Our results showed that the time that pairs spent in close proximity was longer when a dispersing individual was introduced in the home cage. The latency to leave the home cage was longer after the introduction of a dispersing individual. These response variables were not influenced by the relative body sizes of contestants nor by the level of aggression towards each other. In contrast, the aggressive response was significantly influenced by the residency asymmetry established experimentally (“owner” of the home cage vs introduced individual). Our results suggest that the space use by resident individuals is influenced by the dispersal status of conspecifics. The potential ultimate causes driving this effect are discussed.

Seasonal changes in activity and spatial and social relationships of the Iberian rock-lizard Lacerta monticola. - Canadian Journal of Zoology, 79 (11): 1965-1971.

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We conducted a field study to examine the significance of the seasonal changes in activity levels, spatial relationships, and social interactions of the Iberian rock lizard, Lacerta monticola, during the same favorable climatic period. Activity level, movement rate, home-range size, number of same-sex neighbors, and number of agonistic interactions decreased as the mating season finished for males but not for females. The number of females` home ranges overlapped by the home range of each male was also higher in the mating season than in the nonmating season. Measurements of available ambient temperatures showed that the thermal environment was suitable for the activity of lizards in both seasons, therefore the seasonal changes cannot be explained by thermal constraints. Males gain access to several females by increasing their home-range size, thus overlapping a larger number of females` home ranges, but as a consequence, also those of a larger number of male competitors. Thus, the cost of incurring agonistic interactions also increases. These results suggest that because activity and aggressiveness in males are related to their mating success, the balance between the costs and benefits of their activity and spatial strategies differs seasonally, and this may be the cause of the observed seasonal changes.

Aragón, P. & López, P. & Martin, J. (2001) -

Chemosensory discrimination of familiar and unfamiliar conspecifics by lizards: implications of field spatial relationships between males. - Behavioral Ecology and Sociobiology, 50: 128-133.

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The ability of territorial lizards to discriminate between scents of neighbors and non-neighbors might contribute to decreasing the costs of aggressive interactions. To test this hypothesis, we conducted a field study to analyze the spatial relationships between male Iberian rock-lizards, Lacerta monticola. We then used the same individuals in a laboratory experiment to test whether male lizards can use chemical cues to discriminate between familiar conspecific males (those whose home ranges overlapped) and unfamiliar conspecific males (those whose home ranges did not overlap, and whose home range centers were at least 50 m apart). Differences in tongue-flick rates in the presence of chemical cues suggested that male L. monticola discriminated between odors of familiar and unfamiliar males. The behavioral responses were also dependent on relative differences in body size between the responding male and the unfamiliar male that donated the scent: There was a significant negative correlation between tongue-flick rates emitted in cages of unfamiliar males and the body size differences between males. In contrast, when the donor of the scent was a familiar male, the tongue-flick rate was not dependent on body size differences. These results are compatible with individual discrimination through chemical cues in male L. monticola.

We conducted a field study to analyze spatial relationships among male Iberian Rock Lizards, Lacerta monticola. We then used the same individuals in a laboratory experiment to test whether avoidance responses of a male lizard in a previously unknown area is affected by presence of scents of familiar and unfamiliar resident males. Time spent attempting to escape in presence of unfamiliar odors was significantly higher than in presence of familiar odors suggesting avoidance of unfamiliar scent marks, which may lower the costs of aggressive interactions. Our results are concordant with previous studies in which differential tongue-flick rates showed discrimination between odors of familiar and unfamiliar males of L. monticola.

Many studies on contest competition used residency
asymmetry as a discrete variable. However, the
probability of winning an interaction may change as a
continuous function of the value of the location where the
encounter occurs. We performed a field study to examine
the importance of location within a home range and
relative body size to the outcomes of agonistic interactions
between male lizards, Lacerta monticola. The distances to
activity centers (themost used locations based on a density
function of sightings) and relative size play important
roles in agonistic interactions and had interacting effects in
natural conditions. On the other hand, previous studies
with lizards suggested that inferior competitors are able to
avoid agonistic interactions in the field. Thus, we staged
encounters in the laboratory to examine the behavioral
responses of smaller individuals. The responses of each
focal smaller male were measured in its own home cage
(resident), in the cage of a larger male (intruder) and in a
cage in which no male was previously present (control).
The predominant behavioral tactics of smaller males were
avoidance when they are the intruders and displaying
when they are the residents. Submissive displays by
smaller males may help reduce the costs of agonistic
encounters.

Variation in environmental factors plays a central role on organisms` physiological changes. However, the physiological response to predation risk has rarely been investigated in reptiles. Chemical senses are important for intraspecific communication in squamate reptiles. In male lizards Iberolacerta cyreni the maintenance of relative proportions of lipids in femoral gland secretions is costly, which may ensure honest signalling of quality. We hypothesized that increased predation risk should compromise the maintenance of such lipid proportions, as both a fear response and escaping behavior can have physiological consequences. We simulated predator attacks and found that relative proportions of lipids in femoral gland secretions changed in disturbed lizards but not in control ones. Thus, predator–prey interactions may modulate relative concentrations of chemicals in scents of lizards. Potential consequences of this effect on intraspecific chemical communication are suggested.

Aragón, P. & Meylan, S. & Clobert, J. (2006) -

Dispersal status-dependent response to the social environment in the Common Lizard, Lacerta vivipara. - Functional Ecology, 20: 900-907.

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1. Individuals following different strategies such as philopatry or dispersal may also differ in other phenotypic traits, since dispersing individuals have to face novel physical and social environments. There is growing evidence of the use of information obtained from conspecifics in a variety of contexts. It has been demonstrated that before natal dispersal, juveniles of Lacerta vivipara use social information through conspecific chemical cues, and that various phenotypes use this information differently. We hypothesized that, after dispersal, the behavioural responses of yearlings to different social environments assessed through conspecific odours depend on the dispersal status.
2. We tested the response of philopatric and dispersing yearlings of L. vivipara to different types of social cues, controlling for the prenatal and postnatal environment. Each yearling was faced with environments with no conspecific odours, with scentmarks from one or three yearlings that were held isolated during captivity, and from three socially housed yearlings. Thus, we examined the response to the number of donors and to the social environment experienced by donors. We recorded the time spent walking and attempting to escape as indicators of activity and avoidance response, respectively.
3. Philopatric and dispersing individuals reacted differently to the social environments presented through odour marks. This dispersal status-dependent response was not modulated by the prenatal and postnatal factors examined.

Arakelyaan, M. (2007) -

The ratio of blood parasites between hybrids and their parental species of rock lizards of genus Darevskia. - 14th European Congress of Herpetology and SEH Ordinary General Meeting, 19-23 September 2007, Porto, Portugal. p. 170.

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Arakelyan, M. (2001) -

Development of ovaries in bisexual and parthenogenetic rock lizards of genus Darevskia. - In: Ananjeva, N.B. (Ed.): The Problem of Herpetology. Proceedings of the 1st Meeting of the Nikolsky Herpetological Society, Moscow, 21-22. (in russisch)

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Arakelyan, M. (2002) -

The study of age, growth, and longevity in the triploid hybrids of rock lizards of the genus Darevskia in Armenia. - Russian Journal of Herpetology, Moscow, 9 (1): 63-68.

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Skeletochronological study of rock lizards hybrids Darevskia valentini × D. unisexualis, D. valentini × D. armeniaca, D. nairensis × D. unisexualis, and their parental species, collected from three mixed populations (near village of Kuchak, Lchashen, Lchap — mountainous regions of central Armenia) proved their similar pattern of growth. However, the hybrids are characterized by faster rate of growth and predominantly bigger size of body in each age group contrary to their parental species. The longevity of studied hybrids is same as in parental species and reaches 6 – 7 years.

Skeletochronological study of Eremias strauchi (n =18) has shown equable pattern of growth. The lizards under study reach sexual maturity after the 1st or 2nd year of life and have two clutches. Maximum age for the studied sample was 4–5 years.

Age structure of population of the Sand Lizards Lacerta agilis in Armenia. - Program and Abstracts, 12th Ordinary General Meeting Societas Europaea Herpetologica, Saint-Petersburg, Russia, 12–16 August 2003: 31.

The skeletochronological method has been used to assess age distribution and age-related differences in body size among populations of the parthenogenetic lizards Darevskia sapphirina, D. uzzelli, D. armeniaca and D. unisexualis from Turkey and Armenia. The age distribution between D. armeniaca and D. unisexualis did not significantly differ and ranged from 1 to 8 years. Maximum age for both D. uzzelli and D. sapphirina was 6 years, and 8 years for both D. unisexualis and D. armeniaca. In all the studied species, individuals reached sexual maturity after third hibernation. According to patterns of growth marks resorption, D. sapphirina is distinguished from all other rock lizards of genus Darevskia by a narrowest periosteal bone as result of high rate of endosteal resorption resulting in complete destruction of hatchling line and line of the first hibernation.

Morphological and skeletochronological studies on four parthenogenetic species of genus Darevskia. - Abstracts of the 7th International Symposium on the Lacertids of the Mediterranean Basin, 06-09 September 2010, Palma de Mallorca, Balearic Islands, Spain: 62.

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Morphological diversity at 20 pholidotic and 16 meristic characters were studied on three parthenogenetic species Darevskia unisexualis (Darevsky, 1966) D. uzzelli (Darevsky & Danielyan, 1977) and D. sapphirina (Schmidtler, Eiselt & Darevsky, 1994) from Turkey which originated from interspecific hybridization of same parental species D. raddei (Boettger, 1892) and D. valentini (Boettger, 1892) and compared with parthenogenetic species D. armeniaca (Mehely, 1909) which originated from hybridization of D. mixta (Mehely, 1909) and D. valentini (Boettger, 1892) (Murphy et al, 2000, Fu et al, 2000). D. armeniaca significantly differ (ANOVA, Tukey HSD test; p < 0.05) from D. unisexualis and D. sapphirina by 11 pholidotic characters, as well as from D. uzzelli by 9 variables. D. unisexualis has shown significant differences by 8 pholidotic variables from D. sapphrina. Only two pholidotic characters distinguish D. uzzelli from D. sapphirina and D. unisexualis. In addition to meristic characters D. sapphirinia has shown significant less length and height of head as well as length of legs than D. armeniaca, D. unisexualis and D. uzzelli. The average means of snout-vent length of D. sapphirina were less in each age group in comparing with other species. The measurements of bone thickness have revealed the significant high level (p<0.001) of endosteal resorption in femur bones of D. sapphirina removing the first growth layers which usually present in bones of the most Darevskia species. The oldest lizards in our specimens were aged 6 years in D. uzzelli and D. sapphirina and 7 years D. unisexualis and D. armeniaca.

Arakelyan, M. & Stepanyan, I. (2008) -

The size of red cells of hybrids and parental species of lizards of genus Darevskia. - In: Pafilis, P., Kotsakiozi, P. & E.D. Valakos (eds.): 6th International Symposium on the Lacertids of the Mediterranean Basin, 23-27 June 2008, Mythimna, Lesvos, Greece: 15.

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The cytometrical analyses of diploid bisexual species Darevskia
valentini, and closely relative parthenogenetic species D.
unisexualis and D. armeniaca, as well as their triploid hybrids (D.
valentini × D. unisexualis and D. valentini × D. armeniaca) and
tetraploid hybrid D.valentini × D. unisexualis arising from
hybridization in mixed population in vicinity of Kuchak village
(Aragatsotn region, Armenia) has shown significant differences
(P<0.001) in surface and perimeter of erythrocytes between all of
them according to ANOVAs post-hoc test. The size of red blood
cells of triploid hybrids exceeds that of diploids by 25-30%, but
tetraploid male exceeds diploid males by 22% and by 10 % smaller
than those of triploid male hybrids. Among parental species the
smallest parameters of erythrocytes have D. valentini, when D.
unisexualis have the largest red blood cells. Accordingly, the
erythrocytes of hybrids of D.valentini × D. unisexualis were larger
than D. valentini × D. armeniaca.

Arakelyan, M.S. (2001) -

Skeletochronological study of Rock Lizards from Armenia and some questions of their ecolpogy. - Ph.D. Dissertation. Russia, St. Petersburg, Zoological Institute. (in Russisch)

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Arakelyan, M.S. (2007) -

The ratio of blood parasites between hybrids and their parental species of rock lizards of genus Darevskia. - Abstracts, 14th European Congress of Herpetology, 19–23 September, 2007, Porto, Portugal.

Currently among sympatric populations of biparental species of genus Darevskia, which were involved in reticulate evolution,
the processes of hybridization take place only between D. raddei and D. portschinskii species. The successful combination of parental
species with low reproductive isolation may be considered as main factor explaining the origin of parthenogenetic species.

Growth and age composition of some parthenogenetic and bisexual species of Armenian rock lizards (Lacerta). - Entomological Review, 80: 161–166.

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Age determination based on counting the annual layers in the femur was performed for three parthenogenetic species of Armenian rock lizards, Lacerta armeniaca (n = 147), L. unisexualis (n = 68), and L. dahli (n = 18), and for two bisexual species, L. nairensis (n = 29) and L. raddei (n = 32). Comparison of parthenogenetic and bisexual species shows that the growth rate, sexual maturity age, and life longevity in rock lizards are independent of the type of reproduction. The examined Lacerta species reach maturity by the second or third year of life. The maximum age of individuals in the samples studied is 7 and 5 years for L. armeniaca and L. nairensis, respectively, and 6 years for L. unisexualis, L. dahli, and L. raddei.

Range overlap between species even at local scales is a defining trait in the biogeography of the Caucasian rock lizards of the genus Darevskia. Syntopy between populations of two biparental species is remarkable, especially when they are parental contributors of parthenogenetic species. The detailed analyses of such contact zones revealed that when biparental species D. valentini and D. nairensis are in syntopy, no hybrid forms arise. This is likely due to their different reproductive periods. In contrast, the biparental D. raddei and D. portschinskii often produce hybrids, and the parthenogenetic species D. rostombekovi is also found in low numbers in the same sites which suggests that they are an occasional result of hybridization. Three sympatric zones between D. raddei and D. portschinskii are known for Armenia, Nagorno-Karabakh Republic (NKR) and Azerbaijan. Likely, the reproductive isolation between D. raddei and D. portschinskii is not complete which is contributing to reticular evolution. The hybridization events between syntopic D. raddei and D. portschinskii produce lizards with intermediate scalation and coloration characters when compared to allopatric populations. Among 207 lizards from Northern Armenia 59 (28.5%) D. raddei and 53 (25.5%) of D. portschinskii displayed intermediate morphology, 6 (2.86%) were parthenogenetic D. rostombekovi and 4 (1.93%) triploid hybrid females D. raddei × D. rostombekovi. Of 20 lizards with intermediate morphology 53 eggs and 35 young were obtained; 12 of young were similar to D. rostombekovi according to their external morphology. Among 143 lizards collected in another syntopic locality with D. raddei and D. portschinskii in NKR, 24 were hybrid females of uncertain morphological ascription. Among 66 D. portschinskii 18 individuals displayed modified morphological characters similar to D. raddei while among 43 D. raddei 14 individuals show morphologies close to D. portschinskii. Therefore, the successful combination of parental species with low reproductive isolation may be considered as the main factor explaining the origin of parthenogenetic species. For next steps of reticulate evolution, the combination of biparental and parthenogenetic species is required. Namely, in the sympatric zone of biparental D. raddei (40% of lizards) and parthenogenetic D. rostombekovi (35%), D. armeniaca (20%) and D. dahli (5%), triploid hybrids arise only between D. raddei and D. rostombekovi (18 hybrid males and 6 intersexes with both hemipenises and oviducts found). D. raddei from this locality showed pholidosis and coloration approaching D. rostombekovi.

Cytogenetic analysis and the patterns of alloenzyme migration were used to analyse the possible endogamic depression caused by the scarce number of specimens of the lizard still in existence. The karyotype of Gallotia simonyi has a chromosome pattern of 2N = 40 (38 accrocentric and 2 microchromosomes with a pair of NOR located in telomeric position at pair 19). There would appear to be no chromosomic definition for sex nor do variations appear in the layout of C bands over the various individuals. Protein electrophoresis has shown that all individuals are monomorphous over the 32 loci analysed. This fact reveals absence of heterocygosity and polymorphism. From the results, we discuss the possibility of total loss of genetic diversity and the application of other techniques towards confirming this extreme (analysis of mini- and macro-satellites). An optimisation of the present cross breeding system used for the specimens in captivity is proposed in order to palliate the high rates of endogamy.

Climate warming and the decline of amphibians and reptiles in Europe. - J. Biogeogr. 33: 1712-1728.

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Aim We explore the relationship between current European distributions of amphibian and reptile species and observed climate, and project species potential distributions into the future. Potential impacts of climate warming are assessed by quantifying the magnitude and direction of modelled distributional shifts for every species. In particular we ask, first, what proportion of amphibian and reptile species are projected to lose and gain suitable climate space in the future? Secondly, do species projections vary according to taxonomic, spatial or environmental properties? And thirdly, what climate factors might be driving projections of loss or gain in suitable environments for species? Location Europe. Methods Distributions of species are modelled with four species–climate envelope techniques (artificial neural networks, generalized linear models, generalized additive models, and classification tree analyses) and distributions are projected into the future using five climate-change scenarios for 2050. Future projections are made considering two extreme assumptions: species have unlimited dispersal ability and species have no dispersal ability. A novel hybrid approach for combining ensembles of forecasts is then used to group linearly covarying projections into clusters with reduced inter-model variability. Results We show that a great proportion of amphibian and reptile species are projected to expand distributions if dispersal is unlimited. This is because warming in the cooler northern ranges of species creates new opportunities for colonization. If species are unable to disperse, then most species are projected to lose range. Loss of suitable climate space for species is projected to occur mainly in the south-west of Europe, including the Iberian Peninsula, whilst species in the south-east are projected to gain suitable climate. This is because dry conditions in the south-west are projected to increase, approaching the levels found in North Africa, where few amphibian species are able to persist. Main conclusions The impact of increasing temperatures on amphibian and reptile species may be less deleterious than previously postulated; indeed, climate cooling would be more deleterious for the persistence of amphibian and reptile species than warming. The ability of species to cope with climate warming may, however, be offset by projected decreases in the availability of water. This should be particularly true for amphibians. Limited dispersal ability may further increase the vulnerability of amphibians and reptiles to changes in climate.

I visited southern France to search for Timon lepidus for the third time in 1998. Sites visited included (among others) Maussane, Lac du Salagou and the surroundings of this lake, St. Rémy-de-Provence, the surrounding hills of Le petit Crau and Avignon. I did not find Timon lepidus this time, but did see Natrix maura, Malpolon monspessulanus, Lacerta bilineata, Podarcis muralis and many Psammodromus hispanicus. There were not very many amphibians visible because of the hot weather.

The author describes a herpetological vacation in southern France. Reptiles and amphibians were abundant in this region. Areas of interest were: La Petite Crau, La Plaine de Ia Crau, Les Alpilles, Les Rochers de Ia Pene and Le Barrage de Sr. Remy. The author was especially interested in spotting Ocellared Lizards (Timon lepida) bur didn` r succeed in finding them. Nevertheless a lot of other animals were seen, as is shown in the table. Nor mentioned in the table are several Red-eared Sliders (Trachemys scripta elegans), that were seen at Le Barrage de St. Remy. In addition the author visited Le Village de Torrue at Gonfaron (breeding-center for Testudo hermanni and Testudo graeca) and several other pla- ces in the Sr. Remy region.

The author describes a herpetological trip to Portugal organized by a travelling agency specialized in ecological trips. The group consisted ofeighteen people and was especially interested in reptiles and amphibians, though most of the participants were bird-watchers and nature-lovers too. Day by day the author describes the various reptiles and amphibians that were seen. A table is added, in which the num- bers represent the day the respective animals were spotted. The regions that were visited were: Sao Bras {day I), Guadiana river {day 2), Parque Natural da Ria Formosa (day 3), Huelva and Coto de Dofiana in Spain {day 4), Lagoa and the Monchique (day 5), the rocky coast of Carrapateira, Cabo de Sao Vicente (day 6), Almarjao, close to Sao Bras (day 7). Three species of newts and salamanders were seen, eight species of frogs and toads, two species of turt- les, ten species of lizards and five species of snakes. A list was made of the insects, the birds (123 spe- cies) and plants that were seen.
The trip was very successful, with regard to the inte- rest of the group

The Ocellated Lizard is the largest Lacerrid. Males can grow up to 65 cm, females to slighdy less. They are brown to green with a pattem of small black dors (forming rosettes in juveniles) and several rows of blue spots on their sides. Males have very large heads, compared to rhe females. The nominate subspecies inhabits the lberian peninsula, sourhem France and northwestem ltaly. In southeastem Spain the subspecies Timon lepidus nevadensis is recognised. The lizards are found in dry, uncultiva- ted areas, preferably near a large stone or piles of stones.
The author keeps his animals in outdoors enclosu- res for five months during the summer months. The indoor terraria measure at least 1.25x0.60x0. 65 cm (lxwxh) with a 10-15 cm layer of substrate, consisring o f a mixture o f loam, clay, garden soil and marl grit. The terraria are decarared with pie- ces of marl, branches and stubs, attached to the backside, artificial plants and some dry shrubs of for instanee heather, which the animals use to shed their skin. The terraria are illuminated with fluo- rescent lights, including those which produce UV- B light (for insrance Reptisun 5.0). Hearing is pro- vided by an iocandescent spotlight of 80-1 00 W. Passive ventilation is provided by one ventilation- grid near the bottom, close to the spotlight, in con- junction with another grid at the opposite side, near the top. Photoperiod is varied according to the normal cycles at this latitude. The animals stay insi- de from the end of September until mid-May.
The outside enclosures average at 2.0xl.5x0.7 m (lxwxh) and are decorated like the indoor terraria. The pens are covered with plastic-coated wire net- ring, to prevent cats, rats and birds to prey on the animals. This netring is buried in the substrate as well, at a depth of 30 cm, to prevent rhe inhabi- tants ro escape and to keep unwanted burying guests out.
The animals are preferentially kept in pairs; other combinations usually result in aggression and pos- sibly death. Other species (even Lacerta viridis) should nor be kept in combination with these ani- mals.
Their daily activity can start as early as 9.00 a.m and can last to 11.00 p.m.
Ocellared Lizards eat virrually all kinds of inverte- brares, including scorpions and large centipedes. In captivity they will eat all kind of insects, even large beetles. Calcium is ingested by eating marl-grit, water-soluble vitamins A and D3 (Davitamon AD aquosum) are added to the drinking-water, five drops in 50 ml. Riekets used to be a problem, espe- cially with juveniles. But after the animals were given marl-grit and vitamin D3 in the drinking- water, riekets was no Jonger encountered.

During rhe winter, from the middle of December until the beginning of March, rhe spotlights are rurned off and the animals are visible only part of rhe time, some of them even hibernare. No food is given during this period, only water. After several weeks rnaring starts, during the last years ar rhe end of March, and will go on until rhe beginning of May. When rhe female has ro lay her eggs, she digs a tunnel of up to 30 cm length, with a larger hole at the end. Here she lays her eggs. The clutch rypi- cally consists of 5-24 eggs, although in one year the author found two durehes of 26 and 27 eggs from the same female. The eggs are incubated ar 28
(27-31) oewith moist vermiculite or perlire as sub- strate. lncubation can last between 74 and 100 days, but most often between 80 and 95 days.
A table is given of the success-rare during the last eight years. The juveniles are raised in groups, all of the same clutch, and are fed all kind of insects. The receive more vitamins than the adults (5 drops in
10 mi) and marl-grit ad libitum. They are kept warm during the winter.
lt is concluded that Ocellated Lizards are easy to keep and breed.

Activity, temperature and habitat selection of Lacerta monticola were studies at one locality in the Cantabrian mountains. L. monticola has an unimodal pattern and is relatively inactive after midday. There is no significant different in habitat use between males, females and subadults. Lizards do not seasonally vary their patterns of habitat utilization. There is a significant correlation between body temperature and ambient temperature. Also, body temperature is significantly correlated with substrate temperature. Males, females and subadults did not differ significantly in body temperature. Body temperatures varies with time of day but not among months.

Ariëns Kappers, J. (1967) -

The sensory innervation of the pineal organ in the lizard, Lacerta viridis, with remarks on its position in the trend of pineal phylogenetic structural and functional evolution. - Zeitschrift für Zellforschung und mikroskopische Anatomie, Wien, 81 (4): 581-618.

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The sensory innervation of the pineal organ of adult Lacerta viridis has been investigated. Some specimens of Lacerta muralis lillfordi were also used. In the pineal epithelium, a small number of nerve cell pericarya of a sensory type are present. They lie either solitary or in small clusters close to the basement membrane. The axons originating from the nerve cell bodies, i. e. the pineal sensory nerve fibers, first course in the intraepithelial nerve fiber layer which is only locally present and contains a restricted number of unmyelinated fibers. In Lacerta viridis, the pineal fibers generally leave the epithelium at the proximal part of the organ proper. They then form small bundles which run along the outer surface of the basement membrane in the leptomeningeal connective tissue covering. At the proximal end of the pineal stalk the single bundles assemble constituting the pineal nerve. In Lacerta muralis the fibers leave the pineal epithelium at the proximal end of the stalk running farther down within the epithelium. Many fibers become myelinated after leaving the pineal epithelium. The pineal nerve runs ventralward in the midplane just caudal to the habenular commissure to which no fibers are given off. Continuing their ventralward course between the habenular commissure and the rostral end of the posterior commissure which is traversed by some of them, the pineal fibers reach the dorsal border of the subcommissural organ. Small separate aberrant pineal bundles traverse the posterior commissure at various more caudal levels. Having reached the dorsal border of the subcommissural organ, part of the pineal fibers continue their ventralward course directly running along the lateral sides of this organ to reach the periventricular nerve fiber layer lateral and ventral to it. A restricted number of fibers first turns in a caudal direction running between the base of the posterior commissure and the base of the subcommissural organ before turning ventralward to reach the periventricular layer. Most probably, pineal fibers do neither join the posterior commissural system nor innervate the subcommissural organ. Once having reached the periventricular layer, some pineal fibers curve in a rostral direction while others, before doing so, send a collateral in a caudal direction. Both, the main fibers and the collaterals, contribute to the formation of the periventricular layer. The sites of termination of the pineal fibers could not be ascertained.
From the presence of intraepithelial sensory nerve cell bodies and from literature data on the ultrastructure of pineal neurosensory cells it is concluded that the adult pineal organ of Lacerta has a, although rudimentary, (photo)sensory function. The demonstration by our guest-worker Dr. W. B. Quay, of the intraepithelial presence of a tryptamine compound, probably serotonin, points, moreover, to a secretory function of this organ.
In adult Lacerta a well-developed parietal nerve connects the parietal eye with the left lateral habenular nucleus. It traverses the habenular commissure.

A total of 58 (24 MM , 34 VV ) Lacerta viridis specimens, which were collected from Turkish Thrace and the Black Sea region of Anatolia, were evaluated serologically. The analyses support the view that a single race of L. viridis (L. v. meridionalis) inhabits Turkey.

An experimental study of the visual pathways in a reptile (Lacerta vivipara). - Journal of Anatomy, London, 84 (2): 146-167.

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1. A description has been given of the normal histological appearance of thalamic, pretectal and midbrain centres related to the optic tracts in Lacerta vivipara. They are compared with corresponding structures described in other reptiles.
2. Silver impregnation of the brain after removal of an eye has revealed axonal and terminal degenerative changes, essentially similar to those which occur in mammals, except that they develop more slowly.
3. Twenty brains prepared by silver impregnation and two treated by the Marchi method have been utilized for the study of the course and distribution of degenerated fibres of retinal origin. Evidence was obtained for the following main conclusions: (a) That the optic nerve contains only afferent fibres. (b) That decussation of the optic nerves is almost, but not quite, complete. (c) That crossed retinal fibres terminate in the lateral geniculate nucleus, in the nucleus geniculatus pretectalis, in the nucleus lentiformis mesencephali, in the superficial layers of the optic tectum (including the stratum zonale) and in the nucleus opticus tegmenti. A few may also end in the nucleus pretectalis. (d) That the basal optic root is composed entirely of crossed retinal fibres. (e) That uncrossed retinal fibres form a small fascicle on the outer surface of the optic tract, some probably having connexions in the rostral thalamus with the pars ventralis of the lateral geniculate nucleus. (f) That as the optic tract runs on the lateral surface of the thalamus it contains in its deeper part numerous non-retinal fibres. Many of these appear to cross in the ventral supraoptic decussation.
4. The organization of the visual system in Lacerta has been compared with those in Amphibia and mammals. The possibility of homology has been discussed, with special reference to the lateral geniculate nucleus. This work has been carried out under the direction of Prof. F. Goldby of the Department of Anatomy, St Mary`s Hospital Medical School. I should like to express my thanks for the advice and criticism which he has given throughout.

Oman material of four lizard species is described and discussed ; it comprises specimens
of a previously undescribed gecko, Phyllodactylus gallagheri, two lacertids — the poorly
known Lacerta jayakari Boulenger, 1887, and L. cyanura, also described here for the
first time, and a skink, Ablepharus pannonicus Lichtenstein, 1823. These species have no strong affinities with any members of the mainly Saharo-Sindian fauna now occupying
much of Arabia but are closely related to forms occurring to the north of the peninsula.
It is probable that they are relicts of a more temperate fauna which once entered more
extensively into southeastern Arabia. As might be expected, all four species are absent
from the desert areas of Oman and are confined to the disjunct mesic habitats which
exist in the mountains.

The scientific results of the Oman flora and fauna survey 1977 (Dhofar). The reptiles and amphibians of Dhofar, southern Arabia. - Journal of Oman Studies, Special report, 2: 273-332.

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Some 500 Dhofar reptiles and amphibians are discussed and a minimum of 48 terrestrial species recognised for the area of which 5 were previously undescribed; these are a gecko, Hemidactylus lemurinus, and the lacertids Acanthodactylus felicis, A. opheodurus,A. masirae and Mesalina ayunensis; a new subspecies of gecko from north Oman, Bunopus spatalurus hajarensis is also reported. Various other species are reviewed; it appears that Bunopus abudhabi is a synonym of B. blanfordi, which is probably conspecific with B. tuberculatus, Hemidactylus parkeri is regarded as a subspecies of H. turcicus, the Arabian agamas of the Agama cyanogaster group are shown to include 2 species to which the names adramitana and yemenensis are applicable although their status relative to African populations in the group is not clear; A. jayakari is conspecific with A. flavimaculata, and A. neumanni with A. sinaita; Chamaeleo chamaeleon orientalis is similar to more northern populations of C. chamaeleon but C. c. arabicus is very well differentiated and may be a separate species, the use of genitalia in the classification of Acanthodactylus is emphasised and A. arabicus, A. blanfordi and A. schmidti are all given full species status; the recognition of Mesalina as a genus indepemdemt of Eremias is confirmed, it is suggestes that Atractaspis engeddensis may be conspecific with A. microlepidota and that Lytorhynchus gaddi is a synonym of L. diadema; the distinctive population of Echis carinatus in Dhofar appears to be closely related to those in south-west Arabia and adjoining Africa but animals from northern Oman and the rest of south-west Asia are very different. Additional specimens are reported of Bunopus s. spatalurus and Coluber thomasi, both previously known from only 2 individuals. Geographical variations of a number of species is discribed including 3 geckos that show very considerable differences within southern Dhofar: Hemidactylus homocolepis, H. yerburii and Tropiocolotes scorteccii.
Observations on the ecology of many species are noted, especially for the aberrant gecko genus Pristurus, one species of which, P. carteri, behaves like a small, ground-dwelling diurnal agamid. Resource partition, at least amongst the lizards, appears to be largely based on parameters of time, food (especially prey size), hunting method and space. A number of forms occourring in both noreth Oman and Dhofar show differences in niche in the two areas apparently related to the presence of absence of a competitor.
Several species are recorded from Dhofar for the first time including Bunopus spatalurus, Hemidactylus flaviviridis, H. turcicus parkeri, Pristurus minimus, Agama adramitana, Acanthodactylus boskianus,Mabuya tessellata, Leptotyphlops macrorhynchus and Lytorhynchus diadema. The herpetofauna of the area can be divided into a largely northern element of desert-adapted forms widespread in the Arabian peninsula and a more peripheral element in the south which shows considerably more affinity to south-west Arabia than to northern Oman. No reptile of amphibian endemics occur in the forested areas of Dhofar but 3 species present in the drymesic habitats on the north side of the mountains may be restricted to the region.
Acanthodactylus opheodurus, Acanthodactylus felicis, Acanthodactylus masirae, Acanthodactylus boskianus, Acanthodactylus schmidti, Mesalina adramitana, Mesalina ayunensis.

The basic anatomy of the lacertid hemipenis (intromittent organ) and methods for its investigation are described. In many members of the Lacertidae, the hemipenis has a structure quite unlike that of other squamate reptiles: the distal lobes of the retracted organ are complexly folded and there is a well-defined supporting structure of dense connective tissue, the armature. This incorporates blood sinuses and has an intramuscular portion embedded in the m. retractor penis magnus and two club-shaped bodies, the clavulae, that support the lobes in the erect organ. Unarmatured hemipenes occur in some lacertids and, like those of other squamates, possess sac-like lobes in the retracted state, but they are singular in having the lobes invested by the m. retractor penis magnus. It is argued that many of these apparently primitive hemipenes are in fact secondary derivatives of the armatured type.
There is considerable inter-specific variation in hemipenial structure which is described systematically. In some cases this involves differences in size, asymmetry and simplification, which may arise as physical isolating mechanisms and is useful in distinguishing otherwise very similar species, particularly in the genus Mesalina (p. 1253). Other shared derived hemipenial features provide useful information about relationships between species and higher taxa and a summary of the hypotheses that they support is given (p. 1254).

The structure of copulatory organs is used very widely in systematics, both for differentiating species and for working out relationships. Differences between taxa may arise from a variety of sources, including non-homology, differences in other parts of the animal, direct selection on copulatory organs, development of physical isolating mechanisms and pleiotropic events. Physical isolating mechanisms seem likely to account for the abrupt differences, involving size, asymmetry and simplifications, that are useful in distinguishing very similar lacertid species. Although these differences usually seem to arise at the end of a speciation event they can simultaneously be the initiating mechanism in a second one. Copulatory organs appear to have high inherent stability, probably resulting from frequent location in strongly homoeostatic environments, single function, insensitivity to niche shift and inertia due to the need to conform to the genitalia of the opposite sex. This stability may be overridden at times by direct selection on the organs themselves or pleiotropic events. Such changes tend to be retained because efficiency in copulation depends not on any absolute genital architecture but on close conformity of the organs. It is the combination of relative stability and tangible input of varied change, which tends to be retained, that so often makes these structures good indicators of relationship.

Arnold, E.N. (1986) -

A key and annotated check list to the lizards and amphisbaenians of Arabia. - Fauna of Saudi Arabia, 8: 385-435.

Abstract
The Equatorial African group of lacertids comprises nine species which share a number of derived features and appear to constitute a holophyletic assemblage. Revision of the group using morphological characters analyzed by parsimony and compatibility methods results in the following taxonomic changes: Lacerta jacksoni = Adolfus jacksoni, Lacerta echinata = Gastropholis echinata, Bedriagaia tropidopholis = Gastropholis tropidopholis, Bedriagaia moreavi = Gastropholis prasina, the latter species being validated and removed from the synonymy of Gastropholis vittata. Adolfus jacksoni has most features in common with the hypothetical ancestor of the group, while Holaspis guentheri and the species of Gastropholis are the most derived forms. Geographical variation in Adolfus alleni is described and what is known of the ecology of the Equatorial group summarized. Most are forest dwellers with the majority of the more primitive forms living on tree boles and among litter and timber on the forest floor (Adolfus jacksoni, A. africanus, A. vauereselli). Another, Holaspis guentheri, is confined to tree boles but glides between them, while the species of Gastropholis appear to be climbers in flimsy vegetation and Adolfus alleni is ground-dwelling above the tree line. Many of the morphological features characterizing these lizards appear to be functionally related to their particular habitats.

Arnold, E.N. (1989) -

Towards a phylogeny and biogeography of the Lacertidae: relationships within an Old-World family of lizards derived from morphology. - Bulletin of the British Museum (Natural History), Zoology, 55, 209-257.

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Relationships of lacertid lizards were assessed on the basis of 84 primary and 112 binary characters drawn mainly from morphology, including features of the skeleton, external anatomy, various internal soft part systems and two aspects of behaviour. Among features not previously used, or not fully investigated before, are structure of the septomaxilla and nasal passages, arranged of the xiphisternal cartilages, mite pockets, kidney position, ulnar nerve arragement, thoracic fascia, aspects of the hemipenis and its associated muscles, female genitalia and jaw muscles. On the basis of parsimony analysis and compatibilty treatment of this character set, the Lacertidae fall into two main portions: A paraphyletic Palaearctic and Oriental group of primitive forms, from which is derived a holophyletic assemblage of Ethiopian and advanced Saharan and Eurasian taxa.
The former group ist not fully resolvable, but Psammodromus and Gallotia appear to be sister groups and are probably related to Lacerta parva and L. fraasi and then L. brandtii, Podarcis appears to be related successively to L. andreanszkyi, the sister species L. dugesii and L. perspicillata, and perhaps L. danfordi and L. laevis. This assemblage may be related to archaeolacertas and Algyroides. The separation of Lacerta lepida, L. pater and L. princeps from the agilis group, based on chemical evidence, is weakly contradicted by morphology. Takydromus may be most closely related to L. vivipara, and L. jayakari and L. cyanura constitute the most likely sister group of the Ethiopian and advanced Saharo-Eurasian assemblage.
Taxe in the Ethiopian and advanced SaharoEuroasian assemblage form a long essentially pectinate tree with relatively change between the side branches, except for a strong disjunction separating the more primitive from the more advanced taxa. Most of the former fall on two main branches, with ´Lacerta` australis and ´L.` rupicola possibly basal to them. 1. the Equatorial forest group containing Gastropholis, Bedriagaia, ´Lacerta` echinata, Adolfus, ´Lacerta` jacksoni and Holaspis. The first three of these constitute a holophyletic group and the same is probably true of the remainder. 2. Tropidosaura, Poromera and Nucras, the latter being the sister group of the more advanced forms. These include successively the Ethiopian Philochortus, Latastia, Ichnotropis and Heliobolus, Pseuderemias, Meroles and Aporosaura, and Pedioplanis, and then the Saharo-Eurasian Eremias, Acanthodactylus, Mesalina and Ophisops-Cabrita.
It seems probable that the ancestors of modern Lacertidae arose in western Eurasia, where the family is known since the Palaeocene and is still represented there largely by quite primitive forms (89 species and seven nominal genera). The family later invaded Africa, perhaps first in the early or middle Miocene. Relatively primitive lacertids spread widely in largely mesic situations in the Ethiopian region, radiating to some extent (six present genera and 16 species) and producing Nucras and the related series of advaned groups (eight genera and 54 species) whoich show increasing adaptation to xeric environments. These genera tend to have heir most primitive species in the northeast and north of the Ethiopian region. The most advaned gave rise to the Saharo-Eurasian clade, now made up to Eremias, Acanthodactylus, Mesalina and Ophisops-Cabrita. This invaded the arid areas of North Africa and Eurasia, where it is presently represented by 70 species. Many morphological changes in increasingly advanced lacertids may be functionally related to the problems of survival in arid, hot, open environments. Considerable ecological parallelism exists in lacertids, with members of separate stocks occupying similar niches in different geographical areas. Morphological adaptations associated with these niches contribute significantly to the high levels of character homoplasy found in the family. There is also some correlation between the degree of niche differentiation in various groups and the quality of the phylogenies that can be produced from their physical characters. A number of morphological parallels exist between advaned lacertids and New World macroteiids. In the skull at least, advaned lacertids show a complex mixture of paedomorphosis and acceleration.
Nomenclatorial changes are as follows: Cabrita is synonymised with Ophisops, necessitating a new name, Ophisops nictans, for Cabrita jerdonii. Aporosaura is synonymised with Meroles, Platyplacopus with Takydromus, and Bedriagaia with Gastropholis. ´Lacerta` (or Centromastyx) echinata is also transferred to the latter genus and Lacerta jacksoni to Adolfus. ´Lacerta` australis and ´L.` rupicola are put in a new genus, Australolacerta. It is recommended that Lacerta dugesii and L. perspicillata should not be placed in the otherwise very uniform genus Podarcis. Although clearly paraphyletic, Lacerta s. lat. Should be retained at least for the present and, if necessary putative relationships within it indicated by informal groups or subgenera.

Arnold, E.N. (1990) -

Why do Morphological Phylogenies Vary in Quality? An Investigation Based on the Comparative History of Lizard Clades. - Proceedings of the Royal Society of London. Series B, Biological Sciences, 240 (1297): 135-172.

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Phylogenies based on morphology vary considerably in their quality: some are robust and explicit with little conflict in the data set, whereas others are far more tenuous, with much conflict and many possible alternatives. The main primary reasons for untrue or inexplicit morphological phylogenies are: not enough characters developed between branching points, uncertain character polarity, poorly differentiated character states, homoplasy caused by parallelism or reversal, and extinction, which may remove species entirely from consideration and can make originally conflicting data sets misleadingly compatible, increasing congruence at the expense of truth. Extinction differs from other confounding factors in not being apparent either in the data set or in subsequent analysis. One possibility is that variation in the quality of morphological phylogenies has resulted from exposure to different ecological situations. To investigate this, it is necessary to compare the histories of the clades concerned. In the case of explicit morphological phylogenies, ecological and behavioural data can be integrated with them and it may then be possible to decide whether morphological characters are likely to have been elicited by the environments through which the clade has passed. The credibility of such results depends not only on the phylogeny being robust but also on its detailed topology: a pectinate phylogeny will often allow more certain and more explicit statements to be made about historical events. In the case of poor phylogenies, it is not possible to produce detailed histories, but they can be compared with robust phylogenies in the range of ecological situations occupied, and whether they occupy novel situations in comparison with their outgroups. LeQuesne testing can give information about niche homoplasy, and it may also be possible to see if morphological features are functionally associated with ecological parameters, even if the direction of change is unknown. Examination of the robust and explicit phylogeny of the semaphore geckoes (Pristurus) suggests that its quality does stem from a variety of environmental factors. The group has progressed along an ecological continuum, passing through a series of increasingly severe niches that appear to have elicited many morphological changes. The fact that niches are progressively filled reduces the likelihood of species reinvading a previous one with related character reversal. Because the niches of advanced Pristurus are virtually unique within the Gekkonidae the morphological changes produced are also very rare and therefore easy to polarize. Ecological changes on the main stem of the phylogeny are abrupt and associated character states consequently well differentiated. The small geographical distribution of the group reduces the chance of different forms entering the same niche and developing morphological parallelisms. Many species are represented because there are many refuge areas within the geographical distribution of Pristurus and also because the fact that niches are well differentiated reduces the chance of extinction by species competition. Examination of ten other lizard clades, mainly in the Lacertidae, shows that the historical features noted in Pristurus also occur in other groups with robust and explicit morphological phylogenies, such as Meroles and Gastropholis. On the other hand, they are often absent in assemblages where phylogenetic hypotheses based on anatomy are poorly supported or equivocal, for example Podarcis and Pedioplanis. The amount of alteration of genital features is not as closely associated with environmental factors as in many other morphological characters, possibly because ecological shifts do not affect them so directly and because they have their own special causes of change. This partial independence from the environmental factors that appear to control many characters means that genital features may enable some phylogeny reconstruction to be done, even when other systems are unhelpful. There is no evidence among the studied clades that groups with poor morphological phylogenies have inherent general constraints on the development and maintenance of new structural features. It seems that the quality of morphological phylogenies may give some indication of the kinds of history groups have had. Clearly, if the variety of ways in which evolution proceeds is to be understood, groups with both good and bad morphological phylogenies must be investigated.

Phylogenetic analyses, using parsimony and compatibility methods, were carried out on the South African lacertid lizards assigned in recent times to Aporosaura, Meroles and Pedioplanis. These were based on 80 primary and 102 binary morphological characters which were drawn from osteology, external features, muscles, kidneys and reproductive systems. Contrary to some previous interpretations, there are two well-defined clades: Meroles plus Aporosaura, and Pedioplanis; these form successive branches on the main stem of the phylogeny of advanced lacertids. The clades show considerable parallel development of derived features, presumably because they had very similar initial genetic potential. Relationships within the two groups are shown on p. 800 and p. 802. As Aporosaura anchietae is sister taxon to a clade consisting of three of the seven species of Meroles, it has been transferred to that genus. Relationships in Meroles-Aporosaura are very well substantiated, in contrast to the situation in Pedioplanis. This difference appears to be related to the different kinds of evolutionary history that the two groups have had. The Meroles-Aporosaura clade has spread progressively into increasingly stringent and singular aeolian sand environments which have elicited the production of many, often unique, derived character states related to the functional problems of survival in such situations. As these states are rarely duplicated in outgroups, the characters concerned are easily polarized. This, together with their abundance, means that a robust basis for phylogenetic inference is available. In contrast, Pedioplanis exhibits relatively limited ecological radiation of a kind that also occurs in related groups, and the functionally related derived states elicited are fewer and less distinctive. In fact, production of a phylogeny for Pedioplanis is very dependent on genital characters which seem to be substantially independent of the main ecological changes that have occurred in the genus. The premaxilla is embraced dorsally by the anterior processes of the maxillae in most lacertids, but the processes are less extensive in two sister species of Pedioplanis, P. burchelli and P. laticeps. This modified condition also occurs in the genera Eremias, Acanthodactylus, Mesalina and Ophisops, which together constitute a clade that forms the sister group of Pedioplanis. The modification provides extra evidence for the holophyly of the clade, even though presence in some Pedioplanis shows it to be homoplasious.

Estimates of phylogeny may allow historical events to be reconstructed even without a fossil record. The reliability of such interpretations depends not only on the robustness of the phylogeny but also on its topology. Changes in individual features can be traced and general histories of groups developed and compared with each other. Results are ofter surprising, for Instance the sophisticated tail shedding mechanism of lizards turns out to be a primitive feature that has been lost many times. Similarly, ecological analogues may have developed their common characaters in quite different orders. Phylogenies also provide a way of recognizing constraints and the effects of history on present ecological and behavioural patterns.
When using anatomical characters the quality of the apparent phylogenies produced may be related to ecological history: expansion of a group along an ecological continuum into increasingly demanding niches ina small geographical area tends to produce a robust phylogeny (for instance in Meroles), while this is often not so for widely distributed groups that occupy a more modest range of niches (such as Pedioplanis and Podarcis). Non-morphological data may not show this tendency, but can have their own problems.
Lacertids can be referred to clade with many successive branches in Africa and the Saharo-Eurasian arid zones and a less resolved probably paraphyletic complex of more primitive forms in the Mediterranean and wider Palaearctic areas. The African-Eurasian clade shows a general trend towards ground-dwelling and increasingly arid habitats but is ecologically variied. These animals are important to the study of Mediterranean forms because they provide eco-morphological parallels to them (for instance to Algyroides, Psammodromus and the archaeolacertas) and help form a basis for testing hypotheses about function.
Although it is possible to recognise a number of distinct clades among the Mediterranean and Eurasian forms, relationships within and between these are often much less well substantiated. If results are available in time, current work on mitochondrial DNA sequencing will be discussed.

Arnold, E.N. (1994) -

Do Ecological Analogues Assemble their Common Features in the Same Order? An Investigation of Regularities in Evolution, Using Sand-Dwelling Lizards as Examples. - Philosophical Transactions: Biological Sciences, 344 (1309): 277-290.

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Ecological analogues provide an opportunity to assess the regularity of the evolutionary process and some of the factors that control this. By using cladistic methods of phylogeny reconstruction, it is possible to see if two or more stocks have responded in the same way to similar environments, or whether factors such as accident, contingency and previous history have restricted such orderliness. A method is described for comparing the sequences in which ecological analogues assemble their common traits, including ways of handling incomplete data and of testing the statistical significance of the results. The method is applied to situations that mainly involve lizard groups independently associated with aeolian sand habitats. A case where the lineages concerned are closely related is contrasted with two where this is not so. The first instance involves an intrafamilial comparison of three members of the lacertid genera Meroles and Acanthodactylus, which show strong concordance in trait order and great similarity in their independently acquired traits. The other comparisons are of members of different families: Meroles anchietae (Lacertidae), Uma (Phrynosomatidae), Phrynocephalus arabicus (Agamidae) and Pristurus carteri (Gekkonidae). Here concordance in trait order is much lower and independently developed traits often show substantial differences in the various groups. The principal reason for the disparity in results appears to be the much longer and more varied separate histories of the lineages involved in the interfamilial comparisons before they finally entered aeolian sand. These historical differences result in particular independently acquired features developing much earlier in some lineages than others and in the development of phylogenetic constraints and proclivities that influence the detailed ways some environmental problems are solved. Finally, no evidence could be found that traits which evolve in similar sequences in different lineages are developmentally interconnected.

Arnold, E.N. (1995) -

Identifying the effects of history on adaptation: origins of different sand-diving techniques in lizards. - Journal of Zoology, 235 (3): 351-388.

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The effects of previous history on adaptation can be studied in situations where several taxa have been independently exposed to a well-specified problem posed by their selective regimes. This approach was applied to the ways surface-dwelling dune lizards dive rapidly into loose sand when evading predators. Lacertids of the genus Meroles use all limbs and serpentine body and tail movements, behaviour which arose directly in the context of predator evasion in sandy habitats. The agamids, Phrynocephalus and Agama etoshae employ lateral body oscillations and descend vertically, an aberrant method resulting from previous evolution of a blunt snout. The scincid, Scincus and gerrhosaurid, Angolosaurus, both use high amplitude sinusoidal movements of the body and tail, which apparently evolved in a common ancestor of these taxa long before being co-opted to use in sand. In contrast, the scincid, Mabuya acutilabris, holds the body straight, a method that probably originated in the context of concealment at the end of activity.
In the phrynosomatid, Uma, the head oscillates up to 60 times per second, the forelimbs are folded back and hindlimb thrust is supplemented by scales on the upper arms acting like ratchets. Some components of this method originated in the context of slow concealment in firm substrata at the end of activity periods, a basic pattern later co-opted for rapid diving in compact sand with modification of the snout to form a drill. Finally, this elaborated mechanism was transferred to loose sand with further behavioural and morphological change. Rapid sand burial by Phrynosoma species has the same origin as that in Uma but, like Phrynocephalus, includes a strong vertical component. It too is associated with a blunt snout which, however, arose after initial burial behaviour developed, not before.
These results suggest that, once related lineages have diverged, differences may still accumulate even in similar situations. Sand diving also exhibits striking cases of convergence and provides an example of how function can be used to polarize characters, suggest the situations in which derived states arose and indicate their likely order of change.

The east Asian lacertid lizard genus, Takydromus, is well-defined but its relationships to other primitive Palaearctic lacertids are still uncertain and, although it has similarities to the lower Miocene Miolacerta. Takydromus does not appear to be identical with this. The 16 or 17 recognizable species are reviewed and a key provided for their identification; the remaining area of uncertainty about species boundaries is the Takydromus sexlineatus group in Assam. A phylogeny for Takydromus is estimated using a data set of 35 morphological characters drawn from external features, skeleton and soft-parts. There are two main constituent clades which may be informally recognized as Takydromus s.s. and Platyplacopus. Platyplacopus is subtropical and perhaps relict, whereas Takydromus ranges from temperature to tropical areas. The two clades have different ranges of eggs per clutch. In both of them, elongate forms that climb-extensively in flimsy vegetation have evolved and have independently acquired similar features that confer performance advantages in this situation. Interestingly, the order of assembly of these features is markedly different.

Arnold, E.N. (1997) -

Interrelationships and evolution of the east Asian grass lizards, Takydromus (Squamata: Lacertidae): a preliminary survey. - Bulletin of the British Museum (Natural History), Zoology, 64: 63-90.

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The east Asian lacertid lizard genus,Takydromus, is well-defined but its relationships to other primitive Palaearctic lacertids are still uncertain and, although it has similarities to the lower Miocene Miolacerta, Takydromus does not appear to be identical with this. The 16 or 17 recognizable species are reviewed and a key provided for their identification; the remaining area of uncertainty about species boundaries is the Takydromus sexlineatus group in Assam. A phylogeny forTakydromusis estimated using a data set of 35 morphological characters drawn from external features, skeleton and soft-parts. There are two main constituent clades which may be informally recognized asTakydromus s.s.and Platyplacopus. Platyplacopusis subtropical and perhaps relict, whereasTakydromus ranges from temperate to tropical areas. The two clades have different ranges of eggs per clutch. In both of them, elongate forms that climb extensively in flimsy vegetation have evolved and have independently acquired similar features that confer performance advantages in this situation. Interestingly, the order of assembly of these features is markedly different.

The gender of Podarcis and the virtues of stability, a reply to W. Böhme. - Bonner zoologische Beiträge, Bonn, 49 (1-4): 71-74.

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It has recently been suggested that the gender of Podarcis (the European and Northwest African Wall lizards) is masculine, which would change the accepted endings of the names of six species and additional subspecies. This course is rejected, on the basis of the invalidity of the arguments put forward for masculine gender, its potential disturbance of the settled nomenclature and the confusion it would cause among non-specialist users.

Arnold, E.N. (2001) -

Overview of morphological evolution and radiation in the Lacertidae. - In: Pérez-Mellado, V. (ed.): Abstracts of the 4th International Symposium on the Lacertids of the Mediterranean Basin, 7-11 May 2001, Mahon, Menorca, Spain: 10.

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Arnold, E.N. (2002) -

History and function of scale microornamentation in lacertid lizards. - Journal of Morphology, 252 (2): 145-169.

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Differences in surface structure (ober- hautchen) of body scales of lacertid lizards involve cell size, shape and surface profile, presence or absence of fine pitting, form of cell margins, and the occurrence of longitudinal ridges and pustular projections. Phylogenetic information indicates that the primitive pattern involved narrow strap-shaped cells, with low posteriorly overlapping edges and relatively smooth surfaces. Deviations from this condition produce a more sculptured surface and have developed many times, although subsequent overt reversals are uncommon. Like variations in scale shape, different patterns of dorsal body microornamentation appear to confer different and conflicting performance advantages. The primitive pattern may reduce friction during locomotion and also enhances dirt shedding, especially in ground-dwelling forms from moist habitats. However, this smooth microornamentation generates shine that may compromise cryptic coloration, especially when scales are large. Many derived features show correlation with such large scales and appear to suppress shine. They occur most frequently in forms from dry habitats or forms that climb in vegetation away from the ground, situations where dirt adhesion is less of a problem. Microornamentation differences involving other parts of the body and other squamate groups tend to corroborate this functional interpretation. Microornamentation features can develop on lineages in different orders and appear to act additively in reducing shine. In some cases different combinations may be optimal solutions in particular environments, but lineage effects, such as limited reversibility and different developmental proclivities, may also be important in their genesis. The fine pits often found on cell surfaces are unconnected with shine reduction, as they are smaller than the wavelengths of most visible light.

Arnold, E.N. (2002) -

Holaspis, a lizard that glided by accident: mosaics of cooption and adaptation in a tropical forest lacertid (Reptilia, Lacertidae). - Bulletin of the Natural History Museum, London (Zoology), 68: 155-162.

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Holaspis is the most morphologically apomorphic lacertid taxon with 42 or more derived morphological features arising on its exclusive lineage. Nearly all of these confer advantages in three specialised activities, or ameliorate problems resulting from them. The activities are: climbing on the often vertical open surfaces on tree boles and branches, utilising very narrow crevices in wood and beneath bark, and the ability, unique among lacertids, to glide from tree to tree. Although many of the features related to these activities are likely to result from direct adaptation to the situations concerned, exaptation has been critical in the development of gliding. Two behaviours present in the earliest lacertids have been coopted to this activity: rib spreading associated with basking contributes to an effective aerofoil, and balance control associated with running helps maintain appropriate posture in the air. Features originally developed in the context of crevice use also contribute to the aerofoil and a high surface: weight ratio. So, while natural selection has moulded Holaspis for its present activities, multiple accidents of history have also been important, as they also have in the evolution of bird flight.

The Lacertidae appear to be most closely related to the worm lizards (Amphisbaenia) and then the Teiodea. Living forms fall into four main units, a basal Gallotiinae, and the Lacertinae, Tykadromus, and Eremiainae. Relationships among these and available fossils suggest lacertids arose in Europe, where they have been present for at least 50Myr and then spread independently to East Asia (Takydromus) and the Afrotropical region (Eremiainae). The ancestors of living species probably had a broad structural niche involving some climbing ability in mesic situations. There were later multiple shifts to ground-dwelliung and specialised climbing on open surfaces and in vegetation matrixes, and to xeric habitats.
Gallotiinae includes the west Mediterranean Psammodromus and the Canary Island Gallotia which has produced an adaptive radiation in this archipelago including giant and herbivorous species. The Lacertinae occur mainly in Europe and mesic areas nearby. They appear to have initially spread in the Miocene and speciated rapidly to produce many largely allopatric units that are assigned to Lacerta s. lat. All except L. (Darevskia) of the Caucasus and neighbouring regions have relatively few present species. Three of four units have spread more widely including L. (Lacerta), L. (Timon), Algyroides and Podarcis. These have distinctive ecological features that enable them to coexist with the other forms. Podarcis may have been responsible for the restriction of some small-bodied Lacerta s. lat. And Algyroides to small distributions in specialised habitats. Surviving members of Takydromus probably speciated in the late Miocene. They primitively have many derived morphological and behavioural features functionally associated with climbing in vegetation matrixes but some lineages have become secondarily ground-dwelling. A DNA-based phylogeny shows the group has many anatomical parallelisms and reversals.
The Eremiainae are characterised by a distinctive ulnar nerve pattern and hemipenis structure. The group intially spread into mesic habitats in the Afrotropical region. Forms concerned are Omanosaura (E. Arabia), Australolacerta (S. Africa), the Equatorial group (central Africa), Poromera (W. Africa) and Tropidosaura (S. Africa). The Equatrorial African group includes Adolfus, Gastropholis and Holaspis, the latter having many derived features functionally associated with gliding and others that are exaptations to this activity. One clade derived from mesic Eremiainae shifted into xerioc ground niches and developed many adaptations to these. Within this units, Latastia, Phiolochortus, Heliobolus and Pseuderemias may form a clade centred on northeast Africa, while Ichnotropis, its sister Meroles, and Pedioplanis occur in the south of the continent. One large clade invaded Asia giving rise to Eremias, Ophisops, Acanthodactylus and Mesalina, the last three of which later colonised dry areas of North Africa, sometimes on more than one occasion.
There are several series of ecomorphs in the Lacertidae, each of which has developed between 2 and 8 times, although sometimes to varying extents. They involve adaptations to dense vegetation on the ground, climbing in vegetation matrixes, forest environments, large size, rocky situations, and Aeolian sand. Lineage history and character lability have both been important in determining how lacertids have adapted. The relatively limited ecological and morphological diversity in the group is likely to have been due to restricted opportunity rather than limited ability to evolve.

DNA sequence indicates the Lacertidae contain two subfamilies, Gallotiinae and Lacertinae, the latter comprising two
monophyletic tribes, the Eremiadini of Africa and arid southwest and central Asia, and the Lacertini of Europe, northwest
Africa and southwest and east Asia. Relationships within the 108 species of Lacertini are explored using mtDNA
(291 bp cytochrome b; 329 bp 12S rRNA for 59 nominal species, and reanalysis of the data of Harris et al. 1998, and Fu
2000). The morphology of the tribe is reviewed and 64 of its characters (equivalent to 83 binary ones) also used to assess
relationships. The Lacertini are assigned to 19 monophyletic units of 1 to 27 species, recognised here as the following
genera (contents are indicated in brackets): Algyroides, Anatololacerta gen. nov. (L. danfordi group), Apathya (L. cappadocica
group), Archaeolacerta (L. bedriagae), Dalmatolacerta gen. nov. (L. oxycephala), Darevskia (L. saxicola group),
Dinarolacerta gen. nov. (L. mosorensis), Hellenolacerta gen. nov. (L. graeca), Iberolacerta (L. monticola group), Iranolacerta
gen. nov. (L. brandtii and L. zagrosica), Lacerta s. str. (sand and green lizards, L. agilis group), Parvilacerta gen.
nov. (L. parva and L. fraasii), Phoenicolacerta gen. nov. (L. laevis group), Podarcis (wall lizards), Scelarcis (L. perspicillata),
Takydromus (Asian grass lizards), Teira (L. dugesii), Timon (ocellated lizards, L. lepida group) and Zootoca (L.
vivipara). Both mtDNA and morphology indicate that Lacerta and Timon are sister taxa, and DNA suggests further possible
relationships among genera (Fig. 1, p. 6). Neither DNA nor morphology indicates that the archaeolacertas (sometimes
formalised as Archaeolacerta sens. lat.) form a clade. Instead, they are representatives of an ecomorph associated
with living on rock exposures and using the narrow crevices that these contain.
The Lacertidae probably arose in the European area, with the Gallotiinae later reaching Northwest Africa and the
Canary Islands, and the ancestor of the Eremiadini invading Africa in the mid-Miocene. The Lacertini spread through
much of their present European range and diversified, perhaps largely by repeated vicariance, around 12–16 My ago,
producing the ancestors of the present mainly small-bodied genera, which then underwent often modest speciation. Three
units spread more widely: the Lacerta-Timon clade of large-bodied lizards probably dispersed earliest, followed by Algyroides
and then Podarcis. Overall, European Lacertidae show a pattern of repeated spread, often accompanied by restriction
of previous groups. Expansion of Lacertini may have displaced earlier lacertid lineages from all or much of Europe;
while spread of Podarcis may have restricted many other genera of Lacertini. The earlier expansion of the Lacerta-Timon
clade probably did not have this effect, as difference in adult body size restricted competitive interaction with other
forms. Several invasions of more distant areas also occurred: of East Asia by Takydromus over 10 My ago, and more
recently of northwest Africa by Podarcis, Scelarcis and Timon, and Madeira by Teira.
Relationships within the Eremiadini estimated from both mtDNA, and nDNA differ considerably from those based
on morphology. They indicate relatively mesic forms may have diversified widely across Africa and given rise to at least
three independent invasions of arid habitats. MtDNA also indicates that Lacerta andreanskyi belongs in the Eremiadini
and may occupy a basal position there. It is assigned to a further new genus, Atlantolacerta gen. nov.

Lacerta zagrosica, which inhabits rock exposures at altitudes of over 2000 m in the Zagros mountains, is described more fully, including skeletal, hemipenial and other internal features. It lacks the distinctive features of Omanosaura and other Eremiiniae, and a phylogenetic analysis of morphological data of the Lacertinae, based on 64 characters (equivalens to 83 binary ones), fails to associate it with any of the other taxa. But 620 bp of mitochondrial DNA (fragments of the cyt b and 12S rRNA genes) place L. zagrosica firmly and convincingly with its geographical neighbours, the ground-dwelling Lacerta brandtii, although it shows a divergence of 10 % from this species. Morphological differences from L. brandtii are considerable and may be largely related to the different functional demands of the habitats of the two species.

From descriptive to predictive distribution models: a working example with Iberian amphibians and reptiles. - Frontiers in Zoology, 3:8

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Background: Aim of the study was to identify the conditions under which spatial-environmental
models can be used for the improved understanding of species distributions, under the explicit
criterion of model predictive performance. I constructed distribution models for 17 amphibian and
21 reptile species in Portugal from atlas data and 13 selected ecological variables with stepwise
logistic regression and a geographic information system. Models constructed for Portugal were
extrapolated over Spain and tested against range maps and atlas data.
Results: Descriptive model precision ranged from `fair` to `very good` for 12 species showing a
range border inside Portugal (`edge species`, kappa (k) 0.35–0.89, average 0.57) and was at best
`moderate` for 26 species with a countrywide Portuguese distribution (`non-edge species`, k = 0.03–
0.54, average 0.29). The accuracy of the prediction for Spain was significantly related to the
precision of the descriptive model for the group of edge species and not for the countrywide
species. In the latter group data were consistently better captured with the single variable searcheffort
than by the panel of environmental data.
Conclusion: Atlas data in presence-absence format are often inadequate to model the distribution
of species if the considered area does not include part of the range border. Conversely, distribution
models for edge-species, especially those displaying high precision, may help in the correct
identification of parameters underlying the species range and assist with the informed choice of
conservation measures.

We studied morphological and molecular characters in Podarcis bocagei and P. hispanica lizards in the Ria de Arosa archipelago in coastal Galicia, Spain. Contrasting published information about insular distributions indicated that morphological species identification is problematic. Instead, we identified 145 lizards from 13 islands and several mainland populations by a panel of partially diagnostic nuclear protein loci and through the DNA sequencing of a stretch of 297 base pairs (bp) of the mitochondrial cytochrome-b gene. Correspondence between the molecular identifications was complete, with the exception of two lizards that carried P. hispanica mitochondrial DNA in a P. bocagei nuclear background. The combined results indicate past hybridization events and the oversea dispersal of a female lizard over a distance of about 500 m. Fourteen morphometric and 12 meristic characters were measured in 187 lizards from 15 islands and several mainland reference populations. Multivariate analysis revealed significant differences between sexes and between species. Two island populations for which no molecular data were available were identified as belonging to P. bocagei from morphology. Locally, P. bocagei and P. hispanica have an essentially parapatric distribution, with P. bocagei in the northeast and the islands of the inner Ria, and P. hispanica in the southwest and the islands of the outer Ria. Common patterns for island populations are an increase in absolute size, a decrease in three relative head width parameters, and a lower nuclear genetic heterozygosity than on the mainland. No correspondence was observed between morphological and molecular patterns of intra-specific differentiation. Also, linear distance between populations did not help to explain the results. Because the Ria de Arosa is a flooded river valley, the local bathymetry reflects the order in which the islands have become isolated from the mainland since the sea-level rise started c. 14,000 years ago. However, the temporal order of isolation was not associated with the pattern of population differentiation. The island populations of P. hispanica seem to have diverged over multiple genetic and morphological axes, with no obvious relationship to inter-population distance, coastline history, island ecology, or species habitat preferences. The results indicate that genetic drift is the main force driving in population divergence. Oversea dispersal, that is gene-flow, appears insufficient to counter the morphological differentiation between some island populations. Hypothesis on the biogeographical history of P. bocagei and P. hispanica in western Galicia are discussed.

Lacerta monticola bonnali Lantz, 1927, that inhabits central Pyrenees from 1.800 m to ca. 3.000 m a.s.l. in alpine and subalpine environments, should be considered hereinafter as a full species: L. bonnali stat. nov. due to strong differences in pholidosis, design and coloration, as well as ecological (types of rocks among it lives) and biological features (egg incubation time) that distinguish it both from L. monticola Boulenger, 1905 and L. horvathi Mehely, 1904, their closest geographic relatives, as well as from other Archeolacerta from Europe and Near East.

Morphology and taxonomic revalidation of Lacerta agilis garzoni PALACIOS & CASTROVIEJO, 1975. - Abstracts of the 2nd International Symposium of the Lacertids of the Mediterranean Basin, Quinta de Marin, Algarve: 1.

Iberolacerta bonnali is endemic from the Central Pyrenees, where inhabits from the Arriel Massif in the west, to the Aigüestortes mountains in the east. His range is characterized by the greater size of the mountain massifs which inhabits, and by to present granitic (igneous) rock substrates. By other side, his localities are also characterized by lesser insolations and sun radiations, as corresponds to the pyrenaean parts with greater atlantic influence. I. aranica is endemic from the Mauberme Massif and its spurs. His area appears characterized by greater snow accumulations both in winter and spring. I. aurelioi is endemic from the Montroig, Pica d`Estats and Coma Pedrosa massifs. His area is characterized by the greater insolations and sun radiations, and by the presence of lesser alpine belt surface and absolute heights of the inhabited massifs. Also, he inhabits only paleozoic rocks (mainly cambroordovician quarzites). In respect to other species present in the alpine belt: Lacerta agilis appears in localities characterized by high insolation and sun radiations in respect to other pyrenaean lizards. Zootoca vivipara do not present specific traits as lives in all the pyrenaean range and frequently sympatrically with all the other species here studied. Podarcis muralis appears in the comparatively more outer parts, with greater insolations, from the Pyrenees.

Arribas, O.J. (1999) -

Phylogeny and relationships of the Mountain lizards of Europe and Near East (Archaeolacerta MERTENS, 1921 sensu lato) and their relationships among the Eurasian lacertid radiation. - Russian Journal of Herpetology, 6 (1): 1-22.

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Archaeolacerta s. l., in the currently utilized sense results to be a polyphyletic taxon, composed of three monophyletic and phylogenetically well differentiated taxa: a)-Archaeolacerta (s. str.) Mertens, 1921, which includes three species: A. bedriagae, A. oxycephala, and A. mosorensis, distributed by the northern part of the Central Mediterranean (Corsica, Sardinia, and west Balcanic Peninsula). His most related genera seem to be Teira (his adelphotaxon in our study), Apathya, and Omanosaura. b)-Iberolacerta gen. nov., with two subgenera: Iberolacerta s. str. and Pyrenesaura subgen. nov. includes six species: I. monticola, I. cyreni, I. bonnali, I. aranica, I. aurelioi, and I. horvathi. They are distributed by the mountains of Western Europe (Iberian Peninsula, Pyrenees, Central and Eastern Alps and the north of the Dinaric Chains). His adelphotaxon are the Caucasian and Near East species of Darevskia gen. nov. c)-The third taxa is Darevskia gen. nov., which includes the species of the `L.` saxicola complex besides `L.` derjugini, `L.` praticola, and `L.` chlorogaster. They are distributed by Eastern Europe, Caucasus, and Near East. Other clear relationships among some taxa of the Eurasian Radiation [= `Eurasische linie` from (Mayer and Benyr, 1994)] have been found: Algyroides seems to be the sister group of the species considered of uncertain phylogenetic relationships belonging to the `L.` danfordi-laevis group and of Podarcis. `L.` brandtii seems to be a very primitive species within the Eurasian Radiation. Omanosaura and Apathya appear as sister taxa. `L.` graeca occupies a very basal position in the sister group of Podarcis and relatives (see above). `L.` andreanskyi is the sister species of the Darevskia nov. and Iberolacerta nov. clade. The assimilation of `L.` andreanskyi to Teira is very problematic. Also the `L.` parva group seems to be related to Timon. Some groups of species like `L.` brandtii, `L.` parva-fraasii and `L.` danfordi-laevis probably merit generic rank, but it is necessary a more deep study before to take a decision.

Phylogeny and Relationships of the Mountain Lizards of Europe and Near East (Archaeolacerta Mertens, 1921, sensu lato) and Their Relationships Among the Eurasian Lacertid Radiation - Russian Journal of Herpetology, 6 (1): 1-22.

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Archaeolacerta s. l., in the currently utilized sense results to be a polyphyletic taxon, composed of three monophyletic and phylogenetically well differentiated taxa: a) Archaeolacerta (s. str.) Mertens, 1921, which includes three species: A. bedriagae, A. oxycephala, and A. mosorensis, distributed by the northern part of the Central Mediterranean (Corsica, Sardinia, and west Balcanic Peninsula). His most related genera seem to be Teira (his adelphotaxon in our study), Apathya, and Omanosaura. b) Iberolacerta gen. nov., with two subgenera: Iberolacerta s. str. and Pyrenesaura subgen. nov. includes six species: I. monticola, I. cyreni, I. bonnali, I. aranica, I. aurelioi, and I. horvathi. They are distributed by the mountains of Western Europe (Iberian Peninsula, Pyrenees, Central and Eastern Alps and the north of the Dinaric Chains). His adelphotaxon are the Caucasian and Near East species of Darevskia gen. nov. c) The third taxa is Darevskia gen. nov., which includes the species of the “L.” saxicola complex besides “L.” derjugini, “L.” praticola, and “L.” chlorogaster. They are distributed by Eastern Europe, Caucasus, and Near East. Other clear relationships among some taxa of the Eurasian Radiation [= “Eurasische linie” from (Mayer and Benyr, 1994)] have been found: Algyroides seems to be the sister group of the species considered of uncertain phylogenetic relationships belonging to the “L.” danfordi-laevis group and of Podarcis. “L.” brandtii seems to be a very primitive species within the Eurasian Radiation. Omanosaura and Apathya appear as sister taxa. “L.” graeca occupies a very basal position in the sister group of Podarcis and relatives (see above). “L.” andreanskyi is the sister species of the Darevskia nov. and Iberolacerta nov. clade. The assimilation of “L.” andreanskyi to Teira is very problematic. Also the “L.” parva group seems to be related to Timon. Some groups of species like “L.” brandtii, “L.” parva-fraasii and “L.” danfordi-laevis probably merit generic rank, but it is necessary a more deep study before to take a decision.

The study techniques permit us to check, from a morphological point of view, the existence of three well differentiated species in the Pyrenees: Iberolacerta aranica (Arribas, 1993), Iberolacerta aurelioi (Arribas, 1994) and Iberolacerta bonnali (Lantz 1927). ANOVA detects differences which accumulate aiming populations of different species. Characters with greater discriminative power among taxa are dorsalia, gularia, ventralia and the relative diameter of the tympanic scale. In the canonical analysis, first axis separates central-eastern pyrenaean populations (I. aurelioi) in respect to central pyrenaean ones (I. bonnali and I. aranica). Second axis separates central pyrenaean populations living in a small isolate at the northeast to the Garona river (I. aranica) from the western ones living in the main part of the pyrenean range (I. bonnali). I. aurelioi is characterized by high values ol ventral pattern, gularia, dorsalia and ventralia; and Iow values of masseteric and tympanic diameters, (temporal areas covered only by small tiny scales are not uncommon), generalyzed absence of contact among postocular and parietal scales and a lower number of circumanals and femoral pores. This species can be distinguished unequivocally also by his yellow belly, which is absent in the other pyrenaean species. I. aranica is characterized by greater diameters of the masseteric plate, relatively well developed ventral pattern. low values of dorsalia. ventralia, circumanalia and relatively short fore and hindlimbs. Usually shows a very characteristic scale disposition in temporal area, with three great scales (masseteric. tympanic and a great wedge-shaped intermediate scale). I. bonnali has intermediate pholidotic values among I. aurelioi and I. aranica. The three species show a good discrimination degree among them (superior to 75 %). The best discrimination is for I. aurelioi from which specimens rarely are mistaken with the other two species. Between I. aranica and I. bonnali there is a more imprecise discrimination, although ever superior to 75 % and more marked in males. In respect to the intraespecific variability among species from which more than one massif has been studied. I. aurelioi is the one which shows samples with the best discrimination degree among it (correct classification percentages from 71.42 to 91.6 %). In I. bonnali intracspecific discrimination is lower. Only the males from Bigorre reach the 75 % of correct classification, which suggests that this species is relatively uniform.

A survey on the geogtraphical variation of scalation in the Sand lizard was done by means of both univariate (ANOVA) and multivariate (Canonical discriminant Analysis and cluster of the Mahalanobis distances among centroids). Our results show the existence of two clearly separated groups, in concordance with previous authors. Lacerta agilis garzoni is enough differentiated to by considered a valid subspecies. Lacerta agilis argus is a mere synonym of Lacerta agilis agilis. Lacerta agilis chersonensis despite their overall similarity in pattern and coloration to the so-called Balcanic group belongs to the Caucasic one. Data on the currently known habitat and chorology of Lacerta agilis garzoni are given.

Canonical Discriminant (CDA), ANOVA and ANOSIM analyses were calculated for all recen tly known distribution areas of Iberolacerta cyreni including severtal small and unstudied peripheral populations. The monly differentiated sample is Guadarrama (the nominate subspecies), with very limited overlap in the CDA (correct classification > 70%) and different from nearly all the other samples in ANOSIM. Guadarrama is a recently differentiated but well diagnosable (morpho) subspecies (with lower values of dorsalia, ventralia and greater values of circumanalia). Despite the mtDNA differences of the Béjar specimens, their morphology is largely equivalent to that of I. Cyreni castiliana (Gredos), but clearly differ in their female body elongation (near 1 cm) with shorter limnbs, a possible strategy to increase clutch size. Populations from the Sierras de Avila (Villafranca, Serrota and Paramera) are very similar among them. Villafranca (in males) together with Béjar (in females) are the most connected samples in MST, and the root of the species differentiation from a morphological point of view, once discarded geographical and climatic influence on morphology. All populations except Guadarrama shall be considered as I. c. castiliana by their morphological identity with gredos. These morphological similarities probably are the reflect of extensive gene flow among them, responsible of maintaining their morphology largely euqivalent.

The most ancient report of Lacerta agilis in Pyrenees is given. The reference to its presence in Montlouis corresponds to the book about the nature of the Pyrenees written by Eugen Trutat in 1894, twenty two years before Boulenger (1916) publication, considered to be the oldest (both slope from Ariege and la Cerdanya: data from Ax-les-Thermes and Porté).

In this paper we deal on the ultraviolet color (invisible to us): where we can find it, the capability of animals to see it and the advantages that this color perception offers to them. As the simplest way to detect it is the photography, we describe and review how to photograph the UV, as a result of 15 years of amateur experience, searching
and testing nearly in complete blindness due to the lack of practical information about “how to do it”. We describe the different kinds of photography (chemical and digital); the cameras and objectives suitable (both astronomically expensive ones and cheap options); what are the best characteristics that the objectives should have for this purpose; the films suitable for their use in chemical photography; the different filters (current or discontinued) manufactured along the years; and the subtle combinations among the different materials to obtain pure UV photographs. This
kind of scientific photography is mainly used in forensics, forgery detection, art dermatology and less in Natural History, despite the fact that a great part of animals see this color and use it in important questions of their biology as the social behavior, mate choice or the food search.

Sex dimorphism: Males and females of Iberolacerta martinezricai differ at p < 0.001 in Ventralia (MF), Forelimbs length (M>F), Hindlimb length (M>F), Pileus length (M>F) and at p<0.05 in Collaria (M>F) and Dorsalia (M>F). Batuecan lizards are overall less di-morphic than the three species of Pyrenean rock lizards.
Morphological variability: Variability Index calculated (ΣCV) for I. martinezricai was 6.8 in males and 6.0 in females. Results are very similar (although slightly smaller) to the obtained in the Pyrenean Iberolacerta spp. From two chosen characters, variation coefficient in Gularia falls also between the normal values for the bisexual species and it is a bit greater than in parthenogenetic species; or it is clearly greater as in Dorsalia. Nothing suggests a drastic reduction of the variability in the Batuecan lizard but some characters recall for a relatively small morphologic variability in some characters, in parallel with the lack of mitochondrial variability, perhaps due to its reduced effectives.
Allometry: Both in males and females, the most representative character of growth is the body length (SVL). Most other characters match very closely to this SVL with very similar coefficients: limbs, Parietal, Pileus length and the Anal plate width. Masseteric, Tympanic and Anal length also grow, but not in parallel with SVL. Anal length grows considerably and independently of body size. In the case of fe-males, body limbs have a negative allometry respect to female’s body length and Masseteric, and especially Tympanic plates have a negative allometry respect to the total growth.
Parallel variation in escalation: In males the most related characters of escalation are the two rows of Femoralia, and two characters that have contrary signs: Supraciliar Granula (but not very marked, as coefficients are very small) and specially Circumanalia (that varies with total independence). Also there is a small relationship of Supraciliar Granula with Gularia and even more distant with 4th-toe Lamellae and Circumanalia. In females on one side (first axis), both Supraciliar Granula rows are related to Femoral pores, 4th toe Lamellae, Ventralia and Femoralia; and on the other side (second axis) both sided Femoralia are lonely related with Dorsalia, Collaria and Gularia, and just inversely related to Supraciliar Granula.
Sexual maturation is probably reached when lizards are three years old (in his 4CY). Our smallest gravid female was 56.7 mm, and was probably reproducing in the spring of their 5 CY (when 4 years old). Al-though it is possible that some specimens (especially males, but also some females) reach maturity during 3CY, as it occurs in I. monticola. Growth rate (K) in the first years is 0.145696 ± 0.0633135 (95% confidence). Calculated length at infinite (maximum size if the same rate or growth curve was ap-plied throughout the lifetime of the animal) would be: L∞ = 70.036 ± 13.7598 (95%). This estimate of the asymptotic size (70.036 mm) is very close to the true maximum size found in the total sample of adults (69.31 mm). Growth decreases progressively and the resulting curve is fairly straight, being the growth not far from linear and very similar in both sexes.
Longevity: The biggest male (SVL 68.15 mm) and the biggest female conserved (SVL 68.86 mm) had 7 growth rings (two big and five narrow) in the femur. The two wide rings correspond to the two first years of rapid growth (2CY and 3CY), but when lizards reach sex maturation growth decreases, being latter rings very narrow and even difficult to count. Both specimens had presumably 8 years old (they are 9CY animals: the birth year and 7 growths seasons, and captured just in the beginning of their 9 CY).
Tail autotomy: Iberolacerta martinezricai hatchlings are able to break their tails from birth. Observed hatchlings (1CY) in nature had intact tails. In 2CY, 3CY and 4CY already appear broken and regenerated tails in some specimens. In adults, there are less than a quarter of specimens with apparently unbroken tails, and the others fall in the different categories of regenerated ones (tail tip, middle of the tail or the whole tail from basis) in proportions similar to one quarter. There are no differences between both sexes in the different categories of broken tails, but there are significant differences between juveniles and adults. Females have a slightly big percentage of broken tails from its basis. Bright colored tail in juveniles (bluish in I. martinezricai), is lost during their 2CY. It seems to be an antipredatory strategy. Hatchlings of the Batuecan lizard are able to attract attention on its tail by a striking whirl of its distal part, unlike adults that are unable with their stiffer tail (which can move only in very limited cases of danger or excitement).

We have studied copulation time in Iberolacerta (Pyrenesaura) by identifying scars of recent inter-course (up to one week old) in 174 adult females. The vast majority of copulations are just in spring and only a few in late summer. We have verified the presence of late summer copulations in I. aranica and I. aurelioi (already known for I. bonnali). Not really autumn copulations are known. The period of greatest number of copulations is very similar among the three species. Iberolacerta bonnali is the one with the earliest copulation timespan, followed by I. aurelioi and finally I. aranica with a clear and sig-nificant delay differences from the other two. In I. bonnali although there may be some early inter-courses (end of May in specific locations and depending on the year conditions) copulations occur es-pecially during June (78 % of the newly mated females), being the second week the highest point, and some appear until early July. In I. aurelioi is similar, with numerous copulations throughout June (85 %) and first week of July (76 %). We have detected at least one mating to end of July, and reappearing again in low proportion towards the end of summer (26 %, in one of every four females). In I. aranica, who lives in the area of greatest snowfall, the cycle is delayed and copulations begin significantly later than in the other two species, in mid-June (41 % of the studied females) and end by mid-July (48 %), also significantly later than in the other two species. Also we have been detected a copulation in August (4% of the sample) and two recent copulations in late summer (13%). The testicular cycle in mid-September shows very large testicles but not already formed sperm. The large testes size suggests hormone production that could be responsible of the late-summer copulations outside the normal breeding season.

Tail regeneration in 706 Iberolacerta (Pyrenesaura): 300 I. bonnali, 212 I. aranica and 194 I. aurelioi was studied, distinguishing four situations: 0) intact tail; 1) distal part (point) of the tail regenerated; 2) Tail regenerated from its half part and 3) tail broken and regenerated from its basis. The incidence of autotomy (tail break) in Iberolacerta (Pyrenesaura) runs in adults from a 47.5% up to a 68.3%. In juveniles is sensibly lower and goes from 10.9% to 23.8%.In I. bonnali percentage of autotomized tails is almost identical in both sexes (48% in males and 47.5% in females), whereas in juveniles is only 22.4%, without differences between adults but yes between adults and juveniles. In I. aranica, percentage of autotomized tails is 68.3% in males (fairly high), a 55.3% in females and a 10.9% in immatures. There are not significant differences between males and females (despite to be near to significance) but exist between adults and juveniles. In I. aurelioi males have a break percentage of 54.4%, females 59.8% and juveniles 23.8%, without differences between males and females, and contrary to the other two Pyrenean species, also inexistent between adults and juveniles. Comparing the three species among them, only males differ (the difference arises between I. bonnali and I. aranica), and all the other comparisons among females and juveniles are not significant. Juveniles of I. aurelioi (only any population of I. aranica or rarely and few vivid in I. bonnali) present bluish of greenish tails that do not broke until some days after birth and are able to wave sinuously in order to attract the predators attention upon it. Also, the known and possible predators are reviewed.

Caucasilacerta Harris, Arnold et Thomas, 1998 has been considered as a nomen nudum for the last 18 years. The
main reason for this was the lack of a diagnosis or reference to it. Now, some authors argue that a paragraph in the
same paper could be a valid diagnosis and thus Caucasilacerta might be an available valid name. In the present
manuscript I demonstrate that Caucasilacerta is a nomen nudum by: a) the lack of diagnosis or reference to it accompanying
(sic! mandatory in the ICZN) to the new name; b) the alleged diagnosis (fide Busack et al., 2016) is
102 lines away (two pages, including a figure and its legend) and there is not a reference to it accompanying the
new name; and c) the subject of the alleged diagnosis are “the Rock-dwelling lacertids in the Caucasus” (sic.!),
different from the group to which the new name supposedly refers to, the “L. saxicola group” (sic!). A relationship
between both names that can only be ascertained by a specialist in the group and the concerned geographic area.
As a result of that, Caucasilacerta is a nomen nudum, unavailable, and thus it is not even a Junior Synonym of
Darevskia Arribas, 1997.

It is presented two instances of Lacertid nomenclature in which changes in the species names in func-tion of grammatical gender or number of the honored persons have been suggested. However Interna-tional Code of Zoological Nomenclature only rules to concord in gender the species name with the genus one. Whatever other change, reasonable or not, is an unjustified emendation, as nomenclatural priority is mandatory over grammatical correctness. Similar cases on reptile and amphibian nomenclature are also commented.

The hemipenis of all the Iberolacerta species is studied. Overall morphology is very similar among the different species. Hemipenes are symmetrically bilobed, with the apical section shorter than the basal one (I. galani, I. cyreni, I. monticola monticola, I. horvathi, I. bonnali, I. aranica and I. aurelioi) or subequal or with slightly inverted proportions (I. m. astur and I. martinezricai). All are similarly sized and proportional to the animal’s SVL, except in I. aurelioi (and perhaps to a lesser extent in I. horvathi) that have notably small hemipenes. Concerning the hemipenial microornamentation, all the Iberolacerta s. str. (I. monticola, I. galani, I. martinezricai, I. cyreni and I. horvathi) have hemipenial microornamen-tation of crown-shaped tubercles, more or less elongated with a series or a crown of small spinules in their apex. Among these, isolate odd shaped tubercles can appear, probably corresponding to immature states that do not reach their complete development. The Pyrenean species (subgenus Pyrenesaura) are variable. Iberolacerta bonnali and I. aranica have mainly spiny-like tubercles (among which some crown shaped can appear), whereas I. aurelioi presents crown-shaped microornamentation (with scattered spiny-like, as occur in other Iberolacerta species). Our hypothesis is that crown shaped models are derived (or more mature, following an ontogenetic criterion) in respect to the spiniform ones, but some spiny (or finger-shaped) can be secondary reversals by incomplete maturation. These secondary derived and the original primitive spiny-shaped ones are very difficult if not impossible to distinguish. Phenomena as the heterochronies could be beneath these reversals in maturation.

Busack et al. (2016) concluded that the name Darevskia should be attributed to Arribas (1999) instead
of Arribas (1997), rendering the latter name a junior synonym of Caucasilacerta Harris et al.
(1998). In this paper, we provide a detailed answer to Busack et al. (2016) and argue that Arribas
(1997) fulfilled the publication requirements of the International Code of Zoological Nomenclature
(ICZN) in place in 1997 and, therefore, Darevskia Arribas, 1997 is a valid name. In order to clarify
this issue, a case has been submitted to the ICZN. To avoid unnecessary confusion in Old World
Lacertid taxonomy, we recommend that until the commission rules on this case, Arribas (1997)
should continue to be considered the authority of both Darevskia and Iberolacerta.